rN oy = coo @ Pee AN DS PLANTENTUIN; BOTANIC GARDENS BUITENZORG,JAVA,INDONESIA FLORA MALESIANA BEING AN ILLUSTRATED SYSTEMATIC ACCOUNT OF THE MALAYSIAN FLORA, INCLUDING KEYS FOR DETERMINATION, DIAGNOSTIC DESCRIPTIONS, REFERENCES TO THE LITERATURE:s SYNONYMY: AND DISTRIBUTION, AND NOTES ON THE ECOLOGY OF ITS WILD AND COMMONLY CULTIVATED PLANTS PUBLISHED UNDER THE AUSPICES OF THE BOTANIC GARDENS, BUITENZORG, AND THE AUTHORITY OF THE CENTRAL GOVERNMENT OF INDONESIA, DEPARTMENT OF AGRICULTURE & FISHERIES PREPARED ON AN INTERNATIONAL CO-OPERATIVE BASIS UNDER THE SUPERVISION OF SEVERAL DIRECTORS OF BOTANIC GARDENS, KEEPERS OF HERBARIA AND VARIOUS PROMINENT BOTANISTS FOR THE PROMOTION OF BOTANICAL SCIENCE AND THE CULTURAL ADVANCEMENT OF THE PEOPLES OF SOUTH-EASTERN ASIA TO THE SOUTHWEST PACIFIC REGION SERIES I SPERMATOPHYTA VOLUME 4 PARTE Dt GENERAL EDITOR: Dre. 'G.Gi J. VAN. STEENIS SENIOR BOTANIST / BOTANIC GARDENS / BUITENZORG FOR SALE ONLY » NOT FOR EXCHANGE PUBLISHED BY NOORDHOFF-KOLFF N.V. » BATAVIA 1948 AL, Ag “ARDEN NOTICE Flora Malesiana is designed to represent a concise flora of the Malaysian region. The present part is the first published. It contains the Ist instalment of vol. 4 of series I. The following independent series are planned: Series I. Spermatophyta (flowering plants) . : é ; . ca 15 volumes Series II. Pteridophyta (ferns and fern allies) : j . ca 3 volumes Series III. Bryophyta (mosses and hepatics) . . . . . ca S volumes Series IV. Fungi & Lichenes (fungi and lichens) . : . ca 3 volumes Series V. Algae (algae) ; ; ca 3 volumes Wea Vi COE At the present moment preparations have been made necessary for the start of series I only. As soon as possible the other series will be commenced and directed by special general editors.—The area covered by Flora Malesiana is indicated on the accompanying map by the hatched area. PLAN FOR, THE PUBLICATION OF SERYESm SPERMATOP EY LTA Volume 1. Cyclopaedia of Malaysian botanical collectors and collections, by Mrs M. J. VAN STEENIS-KRUSEMAN. This is a cyclopaedia of Malaysian collections of Phanerogams and Pterido- phytes. It contains over 3000 names of collectors, with short biographies, carefully excerpted itineraries, and literature pertaining to the collections. Indispensable when localizing herbarium sheets of Malaysian plants and inter- preting the scant notes on the labels of older collections. Contains also chapters on methods of collecting in the tropics, hints for travellers, notes on erroneously localized Malaysian collections, &c. — Ready for the press, probably available by the end of 1949. Appr. 600 printed pages. Sample treatment at the end of this copy. Volume 2. Malaysian plant life, by Dr C. G. G. J. VAN STEENIS. This is a second edition, much enlarged, of ‘Maleische Vegetatieschetsen’ (1935) by the same author. It deals with all vegetation types known in Malaysia, as far as described in literature and reports, or known to the author by personally acquired field knowledge, their status and interpretation, their relations, origin, distribution within Malaysia, and importance to mankind. Biological phenomena, both ex- plained or yet unexplained will be briefly touched on. This book will be copiously illustrated. The MS. can be expected to be finished in 1950. Appr. 500 printed pages. Sample treatment at the end of this copy. Volume 3. Malaysian plant geography, by Dr C. G. G. J. VAN STEENIS. This volume consists of two parts. Part one deals with floristic plant geography and contains tables showing the distribution of the ca 2200 recognized indigenous genera of Malaysian phanerogams, compiled from literature and from the Herbarium. Further there are chapters on the history of phytogeographical theories and discussions, a provisional list of the genera with their synonymy, an attempt to divide the Archipelago into phytogeographical districts based on the hitherto known facts of generic distribution, and a discussion of the phytogeographical character of the islands or island groups separately. The MS. is far advanced but not yet ready for the press. Sample treatment at the end of this copy. Part two will deal with the historical plant geography of the Malaysian region. This is only in the initial stage. Volume 4. Flora Malesiana proper. Onwards of vol. 4 the revisions of the families will be printed in the sequence in which they are finished, irrespective of the alliance of the groups concerned. Vol. 4 will be opened by an introductory essay containing chapters on the importance of variability in Malaysian plants, special aberrations with which the Malaysian botanist is confronted in the field and with which he must be acquainted to judge their importance in the often scanty specimens available in the herbaria. A chapter is added on the history of Malaysian phyto- graphy. An annotated list of existing revisions concludes the introduction. PROPOSED CONTENTS. OF VOLUME 4 Pe ETCIACE. . Introduction. . General considerations. . History of descriptive Malaysian botany (by Dr H. C. D. DE WIT). General plan of revisions and hints to collaborators. Keys for identifying Malaysian plants. . Annotated list of former revisions. . Photographs of principal botanical contributors to Malaysian botany. . Systematic revisions of families of Phanerogams, incl. Bignoniaceae, Podo- stemonaceae, Droseraceae, Umbelliferae, Convolvulaceae, Dioscoreaceae, Plumbaginaceae, etc. etc. WONDARWNE MANAGEMENT OF FLORA MALESIANA Flora Malesiana will not be available for purposes of exchange; it is for sale only. Co-operating and collaborating institutions can obtain this flora at a reduced price. For subscribers to a complete series the price will be reduced. For substantial collaborators a special reduction will be fixed individually. General volumes 1—3 will also be sold separately to a limited extent. APPLICATIONS TO" RE DIRECTED VEG N.V. Erven P. Noordhoff, N.V. Noordhoff-Kolff, P.O. Box 39, P.O. Box 103, Groningen, Holland. Batavia-C., Java. and in the Americas to: The Chronica Botanica Co., Book Department Waltham, 54, Massachusetts, U.S.A. SCIENTIFIC COMMUNICATIONS concerning Flora Malesiana should be addressed to Dr C. G. G. J. van Steenis, c/o Rijksherbarium, Nonnensteeg 1, Leiden, Holland. INDEX page ‘ page PCetACede, Wet Pe SL 3. Nyssaceae . . 2” 7 See Acunidiaceaesssir. . §, . 3! Philydraceac:\ -. 2) eee e) Ancisttocladaceae: =. 3. 8 Sarcospermaceae . . ”\ . uumee Aponogetonaceae. . ... 11 Sphenocleaceae . | 7) eee Burtanniacedes st 13 Stackhousiaceae . .. [eee PRINTED IN THE NETHERLANDS BY JOH. ENSCHEDE EN ZONEN » HAARLEM y a0 NaN. Sn y/ ae y y \ wy 7 4] A | Aenorieno % MEMORIAE EORUM QUI SE FLORAE MALESIANAE PERSCRUTANDAE DEDERUNT ET NOBIS DUCES ET EXEMPLA FUERUNT GRATO ANIMO HOC OPUS DEDICANT AUCTORES PREP AGE There are only a few things left in common to the displaced and disjointed inhabit- ants of this Earth; they are the things spiritual. Among those treasures of the mind natural science has come to the fore only in the last three centuries, as a lofty and impartial principle that tends to join people instead of disrupting them. Through war, famine and pestilence the undying fire of science has remained a steady beacon. At the inception of a great work, which shall be the demonstration of the united effort of many workers, it seems meet to remember the function of Science, apart from its benificial or detrimental applications. In these days the adjectives ‘pure’ and ‘applied’ have lost much of their meaning, and the Masters amongst us were the least concerned with this classification. Nevertheless, as long as sentiment, politics, greed and bigotry rule this world, a purely scientific endeavour may become a binding force between individual groups, and maybe, even between nations. At the beginning of this great project we therefore see already a dark shadow cast by this unfortunate era, but rather than to dwell on darker thought let us invoke the light that is cast by those that lived and worked in these regions before us, and let us consider this work as an apotheosis of the ideals formulated by Melchior Treub who, half a century ago, became the initiator of co-ordinated scientific effort in the tropics. His ideals are still ours. To the General Editor, Dr C. G. G. J. van Steenis, we all want to express our gratitude for his initiative, for his boundless energy and, especially, for his faith in this project. To discuss the project of the Flora Malesiana from the technical side would be simply borrowing materials from the scientific collaborators. Rather than to plagiarize it seems fit to leave these matters to the specialist. But as Director of the Scientific Institutes, called ‘’s-Lands Plantentuin’ I may express the satisfaction that our Government, almost a century after the foundation of our Herbarium, after 130 years of effort of these institutes, after the publication of innumerable, chiefly disjointed contributions, has decided to further a unified effort. Amongst the many accusations that our workers have suffered in the last years, some at least have some foundation. We have plowed too deep, we were too few, we did not seek enough contact. But the greatest blight of tropical science has been the lack of continuity. Now we may plow deep, but with many to help. Now we may proceed together and, let us hope, with continuity guaranteed. The great ‘Horn of Plenty’, the cornucopia of our Malaysian Flora, which was opened by van Rheede and by Rumphius shall still flow for a long time. I wish great joy to those that shall have the privilege to examine its contents. (L.G.M.BAAS BECKING) Director of ’s-Lands Plantentuin iv 12. . Systematice plantas suas disponit verus Botanicus; Nec absque ordine easdem enumerat. . Fructificationis principium in theoretica dispositione agnoscit; Nec dispositionem secundum Herbam immutat. . Genera naturalia assumit; Nec Erronea ob speciei notam aberrantem conficit. . Species distinctas tradit; Nec e Varietatibus falsas fingit. Varietates ad species reducit; Nec eas, pari passu, cum speciebus obambulare sinit. Synonyma praestantissima indagat et seligit; Nec acquiescit in quacunque obvia nomenclatura. Differentias characteristicas inquirit; Nec inania nomina specifica praeponit veris. Plantas vagas ad Genera amandare studet; Nec rariores obvias fugitivis oculis adspicit. Descriptiones complectentes differentias essentiales, compendiose sistit; Nec naturalissimam structuram oratorio sermone ebuccinat. . Minimas partes attente scrutatur; Nec ea, quae maxime illustrant, flocci facit. Observationibus ubique plantas illustrat; Nec in vago nomine acquiescit. Oculis propriis quae singularia sunt observat; Nec sua solum, ex Auctoribus, compilat. LINNAEUS, Philosophia botanica [INGRODe Cho N After the appearance of Rumpuius’s Herbarium Amboinense, the result of lifelong research into the botanical treasures of the Malaysian Archipela- go, the first comprehensive work on the flora of these islands was begun by C. L. BLUME, the second Director of the Botanic Gardens at Buitenzorg. His Bijdragen tot de Flora van Nederlandsch Indié (Contributions to the Flora of the Netherlands Indies) consisted of numerous brief botanical diag- noses mostly, however, of Javan species. Shortly after followed his Flora Javae and later Rumphia. None of these books represent a ‘flora’; neither completeness was aimed at nor keys were given. The first design for a flora of the whole of Malay- sia seems to have been drafted by the Swiss bota- nists H. ZOLLINGER and his teacher, A. Morirzi.! I have not succeeded in tracing any further results of their plans. Since the publication? of the Flora van Neder- landsch Indié or Flora Indiae Batavae by F. A. W. Mique. (5 vols, 1854—’60)—which was no ‘flora’ in the present meaning of the word, keys being almost absent—no work has been conceived with the object of covering the Malaysian region. MIQUEL’s work? may be considered as a more or less critical compilation of descriptions, mostly copied or extracted.* MIQUEL must have realized that by his Flora the proper work was only started. This may be concluded from the series of revisions which MIQUEL, together with some specialists, published in 4 volumes Annales musei botanici lugduno batavi (1863-’69),° Choix des plantes rares ou nouvelles (1863), to which was added his posthumous //lu- strations de la Flore del Archipel Indien (1870—71) by his successor at Leyden University, W. F. R. SURINGAR. Unfortunately, Mique had few pupils® which caused a serious shortage of well-trained systema- tists during half a century of botanical endeavour in the East Indies. The only Dutch scientists study- ing the Malaysian flora were: P. DE Boer, who wrote his doctor’s thesis on the subject De Coniferis archipelagi indici (1866), and later became a professor of Pharmacology at Gro- ningen University, and R. H. C. C. SCHEFFER, an extremely able bota- (1) ZoLLINGER, Observationes phytographicae etc. Natuur- & Geneesk. Arch. 1 (1845) 375; ef. also J. K. HAssKARL, Flora 30 (1847) 299. (2) Made possible by a grant of the Ministry for the Colonies. (3) Dates of publication of the several parts in Bull. Jard. Bot. Btzg III, 13 (1934) 284. (4) Compare ZoOLLINGER, Natuurk. Tijdschr. Ned. Ind. 13 (1857) 292-322; id. (in German), Vierteljahrschr. Naturf. Ges. Ziirich 2 (1857) 318-349. (5) Dates of publication of the several parts cf. Nakal, Journ. Arn. Arbor. 6 (1925) 211-213. (6) Cf. the article in honour of Dr A. A. PULLE, who resumed MIQuEL’s work at Utrecht Uni- versity, Bull. Jard. Bot. Btzg III, 16 (1939) 103-105. nist whose thesis was entitled De Myrsinaceis archipelagi indici (1867). SCHEFFER was subsequently appointed as the (fourth) Director of the Botanic Gardens, Buiten- zorg, and ardently promoted the study of the Malaysian Flora, notwithstanding his feeble health. In his term of office he published several important papers, most on Annonaceae’ and Palmae.® De Boer had one pupil in systematic botany, TH. VALETON, who obtained his doctor’s degree on a monographic study of the Olacineae.? He eventu- ally was employed as a bacteriologist in the Sugar Experiment Station in Java but, soon after, joined the staff of the Botanic Gardens, Buitenzorg (1892). After the appointment of Dr M. Treus as the fifth Director of the Gardens in 1880, interest in the promotion of knowledge of the Malaysian flora revived, but TREUB was badly handicapped by the absence of trained Dutch systematists. TREUB—a contemporary of HoOoKER, EICHLER, BENTHAM, and HARVEY & SONDER, the editors of respectively the Flora of British India, the Flora Brasiliensis, the Flora Australiensis, and the Flora of tropical Africa—was well aware that systematic botany in the Netherlands Empire was on the verge of falling behind that in other tropical countries. He judged the advancement of systematics of pre- eminent importance. He engaged W. Burck, a pupil of SURINGAR’s at Leyden, later a teacher of botany at Buitenzorg, as a subdirector of the Gardens (1883) and charged him with critical research into Sapotaceae (getah- pertja family),'° Mucuna,'! the Erythroxylaceae (cocafamily),!2 and Dipterocarpaceae,'* mostly fa- milies of economic importance. TrevuB, who tried continuously to raise a world- wide interest in the Gardens and its botanical in- stitutes, considered the compilation of a new Ma- laysian Flora to be premature. Collections were inadequate and of the vegetation of the surround- ing regions little was known. He advanced, therefore, the idea of composing a local flora of the surroundings of Buitenzorg, covering the region from the mangrove of Tand- jong Priok to the summit of Mt Gedeh at 3000 m. All altitudinal zones would thus be represented. This Flore de Buitenzorg would serve as a guide to botanically interested visitors of the Gardens and be equally acceptable to residents of Java. Dr J. G. BOERLAGE, then conservator of Leyden Her- barium, during a visit to Buitenzorg as a stipen- diate of the Dutch Buitenzorg Fund, had already made collections for the new flora (1889) and published an article on the grasses. '* (7) Ann. Jard. Bot. Btzg 2 (1885) 1-31. (8) Ibid. vol. 1 (1876) 103-164; O. BEcCARI, Relig. ScHEFF. ibid. 2 (1885) 77-171. (9) Critisch overzicht der Olacineae (1886). (10) Ann. Jard. Bot. Btzg 5 (1886) 1-85. (11) Ibid. 11 (1893) 183-190. (12) Ibid. 11 (1893) 190-194. (13) Ibid. 6 (1887) 145-249. (14) Ann. Jard. Bot. Btzg 8 (1890) 47-78. VI FLORA MALESIANA [ser. I, vol. 4! Treus, however, found it difficult to rally workers to this local flora and so most of it was assigned to foreign visitors who sometimes were temporarily employed at the Gardens. Six volumes appeared viz the Myxomycetes by O. PENZzIG (1898), Ferns and Fern Allies by M. RActBorsKI (1898), Hepatics by V. SCHIFFNER (1900), Algae by E. DE WILDEMAN (1900), and Mosses by M. FLEISCHER (1900-22, 4 vols). The 6th and only volume on Phanerogams was written by J. J. SmitH (Orchida- ceae 1905, atlas 1908-’14). None of the volumes of the Flore de Buitenzorg bears the character of a local flora; the majority deal with the whole of Java. FLEISCHER’s Musci even expanded to a standard work on the world’s mosses. Of the flowering plants apart from the Orchida- ceae, much material was collected by BuRCK and H. HALLIER who planned to elaborate a 7th volume of the Flore de Buitenzorg. A list of the species to be included is kept at Buitenzorg, but nothing ever appeared in print. During this period important revisions of fami- lies were published abroad by O. BEccarr in his 3-volume Malesia. Several monographs appeared in the 4° tomes of the Annals of the Royal Botanic Gardens, Calcutta, on the genus Ficus, the oaks and chestnuts, the bamboos, etc. Local floras of other parts of Malaysia were the 3rd edition of BLANcOo’s Flora de Filipinas (1877- °83)! by Naves & F.-VILLAR, SCHUMANN & HOLLRUNG’s Flora von Kaiser Wilhelmsland (1889), and SCHUMANN & LAUTERBACH’s Flora der Deut- schen Schutzgebiete in der Siidsee (1901) with the Nachtrdge (1905). These eastern floras resembled enumerations and were mainly indices of materials collected on expeditions. In 1890 BoERLAGE previously having published two critical studies of Malaysian plants, viz the genus Achyranthes? and the genera of Araliaceae,? started a work of quite another nature in the com- pilation Handleiding tot de kennis der flora van Nederlandsch Indié.+ This comprised a description of the families and genera of Malaysian phanero- gams. The species were—especially in the last parts—only briefly enumerated. He added to a few families keys to the genera. The generic de- scriptions were mostly critically copied from BENTHAM & HOoOoKeErR’s Genera Plantarum, and occasionally emended. Phytographically Boer- LAGE’s Handleiding brought hardly anything new, but now a comprehensive review in the Dutch language of families and genera came within reach of interested persons in the colonies. However, as will be demonstrated later, this interest was and is still more directed towards species than genera. (1) On the dates of publication see MERRILL, Philip. J. Sc. 12 (1917) Bot. 113-117. (2) Ned. Kruidk. Archief II, 5 (1889) 420-430. (3) Ann. Jard. Bot. Btzg 6 (1887) 97. (4) In total 5 parts appeared, the last posthu- mously (1890-1903, 3 vols). The publication was made possible by a grant of the Ministry for the Colonies. BOERLAGE’S work was more intended as a prelude to a general flora than as a final work. He accepted (1896) the post of subdirector of the Botanic Gardens and Head of its first Division (Herbarium and Botanical Museum), as a suc- cessor to BURCK and began a monograph of the Annonaceae>. Unfortunately he soon (1900) fell a victim to a tropical disease while on a tour in the Moluccas attempting to re-collect the plants mentioned by Rumputius in his Herbarium Am- boinense. Another flora was started, at TREUB’s instigation, of trees growing in the island of Java. This was to be based mainly on the collections made by Forest officer S. H. KoorpDErRS who gathered in the field notes on each species (occurrence, value, uses, etc.). Scientific descriptions and keys were by Tu. VALETON. This work is Bijdragen tot de kennis der Boomsoorten van Java (Additamenta ad cognitionem Florae Javanicae, pars 1, Arbores). Thirteen vol- mes compose this standard work, the 12th volume is by J. J. Smirn, the concluding 13th by SmiTH and VALETON. The work was begun in 1894, and finished in 1913. Later illustrations were edited by Koorpers in his unfinished Atlas der Baumarten von Java (4 vols, 1913—18). The Bijdragen is an excellent work with critical descriptions and notes, and still very useful though, of course, now anti- quated. The descriptions of the species and genera are both in Dutch and Latin. During Treus’s directorate many collections, specially of the Outer Provinces,® were brought together. HALLIER made an important one in West Borneo, KoorDeErs in Java and North Celebes, the SARASINS collected in Celebes, ForBEs and Koor- DERS in Sumatra, ForsBes in Timor, while WArR- BURG’S, SCHLECHTER’S, and BECCARI’s great col- lections equalled those of TEYSMANN’s and extended over the whole archipelago. These collections were partly inaccessible though together they could have served to a large measure as a reliable basis for a Flora Malesiana. Lack of trained taxonomists induced TREUB to engage J. J. Smiru, formerly an assistant curator of the Gardens, for taxonomic work. His revisions of Javan Euphorbiaceae, Ulmaceae, Urticaceae, and Orchidaceae proved his ability, and SmirH spent his life in describing Malaysian Orchids, Ericaceae, and Epacridaceae. Unfortunately, he did hardly any monographical work. For the same reasons TREuB selected C. A. BACKER, a teacher in a primary school at Batavia who possessed already a thorough and critical knowledge of the local flora. BACKER intended to fill the still existing /acunae in the phanerogamic part of the Flore de Buitenzorg, which resulted in the publication of one volume of a Flora van Batavia (1907). This was followed by a preliminary schoolflora’ and later by the Schoolflora (1911). (5) Icon. Bogor. 1 (1899) 79-208, t. 26-75. (6) That was: Netherlands Indian territory out- side the islands of Java and Madoera. (7) Voorlooper eener Schoolflora van Java (Pre- cursory Schoolflora of Java). Batavia (1908). Dec. 1948] Introduction VII The latter excellent work contained only + 25 °% of the Javan flora (Choripetalae). He later devoted all his time to the Javan flora, wrote (together with VAN SLOOTEN) a weed flora of tea plantations (1924), 3 instalments of a Handboek voor de Flora van Java (1924—’28), a weed flora of sugar planta- tions (1928-34; vol. II (atlas) not yet completed), and is now engagedincompleting the Flora of Java. ! An ill-advised enterprise was a flora of Java by S. H. Koorpers who, when charged by the N.I. Government to write a flora of the Javan moun- tains, abandoned this concept and hurriedly com- piled an Exkursionsflora von Java (Jena, 1911-12, 3 vols) which did more harm than good and is scarcely of any value to a student of the Javan flora. The flora of the Malay Peninsula was originally included in the Flora of British India, but as the account remained very incomplete KING & GAMBLE, and RIDLEY, started to work on it, pub- lishing a true model of a critical local flora.2 This was later followed by RIDLEy’s decidedly uncritical Flora of the Malay Peninsula (5 vols, 1922-25). On the Flora of Borneo a most helpful Biblio- graphic enumeration of Bornean plants was prepared by Dr E. D. MerRRILL.? In the Philippines MERRILL, after 1902, energetic- ally undertook the research of the Philippine flora, this first resulting in an excellent local Flora of Manila (1912), in a large number of papers dealing with several aspects of the Philippine flora, and crowned by his Enumeration of Philippine flowering plants (1923—’26). The results of frequent expeditions into the Dutch and German territories of New Guinea were published by Dr A. A. PULLE and others in the serial Nova Guinea (vols, 8, 12, 14, and 18), and by C. LAUTERBACH and others,* and in recent years those of Dutch and British parts by MERRILL and other collaborators.> The undesirability of compiling, at this stage, local floras in Malaysia. The studies of the mate- rials of various separate regions persuaded some leading Dutch botanists in the first quarter of our century—for some reasons they doubted the feasi- bility of a Malaysian flora as a whole—to propose several local floras e.g. one of Java, of Borneo, Sumatra, Celebes, etc. This caused the appoint- ment of HALLIER at Leyden to write a Flora of Borneo resulting in a small preliminary paper.® It is clear that this was a wrong policy, born from (1) Seven parts of a mimeographed emergency edition were issued up till now through the care of the Rijksherbarium, Leiden (1940-’48), 9 vols. (2) The contributions of the former appeared under the title Materials towards a Flora of the Malay Peninsula in various numbers of the Journ. Asiat. Soc. Bengal, vol. 58 onwards (18891-915). (3) Journ. Str. Br. Roy. Asiat. Soc. Special number (1921). (4) Under the title Beitrdége zur Flora Papuasiens in many volumes of the Botanische Jahrbiicher (1912 onwards). (5) Journ. Arn. Arb. 9 (1928) et seq. (6) Beih. Bot. Centralbl. 2. Abt. 34 (1916) 19-53. either ignorance of the taxonomic position and the technique of writing revisions, or from the wish for dodging obstacles; the difficulties should be faced directly. Only temporary profit may be gained from making local floras, and both valuable time and money are wasted by the enormous duplication which is unavoidable when the goal of a flora of a plant-geographical unit is to be reached along this tortuous road. The natural sequence is to start with the large flora, eventually followed later by local floras, a procedure followed in the great floras of South America, tropical Africa, India, and Australia. The unnatural sequence of starting with the local flora has led, both in North America and Europe, to a most regrettable state of affairs. The absence of a general flora is also one of the causes that the flora of Java which BACKER has studied close on forty years is only now more or less to be completed. It contains several families which cannot be critically treated (Lauraceae, Araceae, Zingiberaceae, etc.) lacking revisions of these families in the whole Malaysian region. General Flora. A general flora was and is needed and prospects at the end of the first World War seemed favourable. The Forest Research Institute and the Museum for Economic Botany’ at Bui- tenzorg requested much service and urged the Herbarium of the Botanic Gardens to produce speedy results. This induced the Goverment to add to the staff of the Herbarium R. C. BAKHUIZEN VAN DEN BRINK (1917)—he was originally a plantation assistant—Dr D. F. vAN SLOOTEN and Dr H. J. LaM, the first pupils of PULLE at Utrecht (1919). In 1921 Dr H. C. CAMMERLOHER, a German biolo- gist, was appointed, and a professional collector engaged, H. A. B. BUNNEMEIJER. At the same time a scheme was made for critical revisions. These were to be published in the Bul- letin du Jardin Botanique, Buitenzorg® under the heading: Contributions a l’étude de la Flore des Indes Néerlandaises. Economically important fami- lies had priority. The method of treatment stood below that of KING & GAMBLE’s Materials in so far that descriptions were only admitted if species were new or critical. This was believed to save time. On the other hand extensive lists of herbarium numbers had to be compiled. If the latter had been left out and instead a concise characteristic of the occurrence of the species given, besides a good diagnostic description of each species, the Con- tributions would have made a most satisfactory foundation. Though the later Contributions are far more complete than the earlier, the manner of treatment and publication is so laborious and slow that at this rate the Flora Malesiana will never be completed. Till the present 34 Contributions have appeared, comprising 2000 species. Due to the post-war economic depression of 1921—’22 the Staff of the Buitenzorg Herbarium (7) Head of this Museum was the late K. HEYNE, author of the standard work on useful plants of Indonesia (1927). (8) Bull. Jard. Bot. Btzg III, 5 (1923) 294 seg. VII FLORA MALESIANA [ser. I, vol. 4! were reduced, and though towards 1930 there were a few constructive moments, a protracted slump set in after that year and the Staff at Buitenzorg were reduced to the barest minimum. Shortly before the Pacific War the Staff again increased but the circumstances limited advancement of the Flora to planning. I have always felt it as a shortcoming, and not in accordance with the standing of the great work at hand, that the contributions appeared in a peri- odical as scattered articles and not as a separate publication. The work was undertaken on full official au- thority but being printed in an irregularly inter- rupted series of articles in many volumes of a tech- nical journal, it was practically inaccessible to a wider non-professional public. A standard work of this scope and weight meant to be used by future generations and worthy of the wonders of nature in this great land ought to have commanded con- siderable interest in and beyond the tropics, speci- ally so in neighbouring countries. It would not have made a difference in expenditure to issue this work as a separate publication thus materially augmenting its practical importance, its intrinsic value remaining, of course, the same. This seemingly trivial technical-editorial point had very undesirable consequences. If the Govern- ment had once for all decided to order a standard work on the Malaysian flora to be written with all possible expediency and to be used many years afterwards, the halting and haphazard progress in the decade preceding the Pacific War would never have occurred. It is a gratifying thought that the turbulent times of the present could not prevent the Government now to put the Flora Malesiana in an advantageous and satisfactory position both as regards effective publication, and national and international colla- boration of systematists. Co-operation with foreign colleagues, whose help is invited and whose help is needed in order to finish the work within a reasonable time, will now, presumably, more easily be obtained. Evidently, it is far more attractive and stimulating to be entrusted with an individual part of a standard work than with writing an article in a journal. Prospect and scope of the Flora Malesiana. A general flora of Malaysia must result from a careful study of all previous publications, blending them into a harmonious whole, and so founding Malay- sian botany on a secure base of historical fact, ob- servation, and accurate description. This is, how- ever, the labour of a lifetime, and although I may be privileged in witnessing the laying of the founda- tions and the issue ofanumber of volumes, I cannot hope to bring it to a conclusion; progress, more- over, will depend entirely upon circumstances at present beyond control. I have no doubt that when I will be called to abandon this endeavour the historical necessity for the completion of this work will compel someone to continue this task and, eventually, to finish it. It would, however, be wrong were I to convey the impression that this arduous undertaking had entirely originated with myself: on the contrary during many years the conviction has grown among plant taxonomists that the ample collections ac- cumulating in this country warranted the prepa- ration and publication of a Flora Malesiana. The collections are undeniably extensive having been gathered over a wide extent of country.! As I am anxious to render each portion of the work in itself as complete as possible, and desirous of enlisting those of our fellow-botanists as may be willing to take care of those families or groups they are most familiar with, the Flora Malesiana, when terminated will probably consist of a series of local-monographs. For these reasons it seems inadvisable and most inconvenient to arrange the families in the mode of sequence usually adopted in systematic works. I consider it important that the Flora Malesiana should embrace as wide an area as possible, being firmly convinced that no species can be properly defined, until it has been examined in all variations induced by the differences in climate, locality, and soil, which an extensive area affords. Also, the flora of an area cannot be worked out thoroughly with- out a knowledge of the botany of the surrounding countries (these have many plants in common), and so the greater the area encompassed, the better it will illustrate habits, forms, and variations of the species comprised within it. For this reason we have extended the limits of our Flora from Sumatra to New Guinea and from Luzon to Christmas Island, Timor and New Guinea. The use of the Flora Malesiana. In the preceding pages I have mentioned several times the public and the government. Both have a right to a clear understanding of the use of a flora of the scope and character of that now contemplated. Although it is difficult to explain theoretically the ‘use’, i.e. the material benefit of purely scien- tific standard works, many anecdotes and instances concerning scientists entirely possessed by their inventions, instruments, and desire for research, told in biographies and popular literature, exem- plify the eminently practical results based on seemingly impractical and abstract study. The same can be said about this Flora. Botany is not a cherished source of pleasure and interest to naturalists only; and I have but vague ideas of (1) Collections have increased enormously. From 1917 on, the Forest Research Station at Buitenzorg accumulated materials of arboreous plants from the islands outside Java (more than 30.000 numbers): The Museum for Economic Bo- tany furnished by its own collectors another 6000 numbers of those islands. The collectors of the Buitenzorg Herbarium in the past 30 years added to the collections more than 125.000 numbers. A similar increase of Malaysian collections in these last decades is due to the activities at Manila and Singapore; besides, private collectors substantially augmented the collections of New Guinea. A conservative estimate of the collections at Buiten- zorg alone runs to about 400.000 numbers of Malaysian plants. Dec. 1948] Introduction IX possible advantage and ultimate gain for the com- munity and practice by means of this registration of the Malaysian flora. I could refer, of course, to the fact that all other civilized nations have already made considerable progress in the task of making common knowledge of their vegetable resources. Actually the disentangling of confused species, the description of new or the rehabilitation of obsolete genera, the dissection of dried flowers and, in general, the establishment of law and order in ‘the hay loft’, and the publication of the results haveless appeal to the lay public than the segregation of a new promising variety of rice or sugarcane, or devising a method to suppress a pest of coffee or of coconut plantations. The Flora of Malaysia contains besides highly interesting and even unique plant forms, instruc- tive vegetation types, and peculiar ecological and phytogeographical problems, numerous important industrial plants and economic products which, in their manifold kinds, add to human comfort and social prosperity, while, in their ranks, many treas- ures still await discovery, the latest accessions being pectin and mannan producing plants. Their value has come as a surprise both to taxonomists and economists. Nearly a century ago, one of the foremost of British botanists, Sir JosEpH DALTON HooKeErR! wrote an introductory essay to the Flora of British India, one of the most instructive general essays ever written on tropical botany. This nearly one century old exposition of facts and thought meets the present state of knowledge of the Malaysian flora admirably. Its excellence induced me to copy the following from it:— “With regard to economic botany, it is obviously impossible to do more than briefly enumerate, under their respective species, the various products which have been used in the arts: for detailed ac- counts of their value, we must refer our readers to the many excellent works on those subjects, which have been published by Indian botanists.” “Our work is intended to facilitate the progress of economists, by supplying their great desidera- tum, a critical description of the plants which yield the products they seek. We have had a considerable experience both in medical and economic botany and we announce boldly our conviction, that, so far as India is concerned, these departments are at a standstill, for want of an accurate scientific guide to the flora of that country. Hundreds of valuable products are quite unknown to science, while of most of the others the plants are known only to the professed botanists. The mass must indeed always remain so: just as the refinements of the laboratory and the calculations of the mathema- ticlan must ever be mysteries to the majority of manufacturers and navigators, whose operations are based on the sciences in question. It is a mistake to suppose that it can be otherwise; or that those who are engaged in forwarding a science so exten- (1) Hooker & THomson, Flora Indica (1855) 1-280, specially p. 3 et seq. sive and abstruse as philosophical botany, can command the time to become so familiar with the details of the commercial value of vegetable prod- ucts, as to be safe referees on these subjects. On the other hand, it is equally a mistake to suppose that those who devote themselves to the collection of economic products, can possess the experience and botanical knowledge necessary to render their identifications of tropical plants trustworthy in the eyes of men of science. It is therefore as a strictly scientific work that we offer this commencement of the Flora Indica to the public, but though the advancement of abstract science is indeed its pri- mary object, yet as we yield to none in our estimate of the value of economic botany, we confidently trust that... our labours will be found of material service.” “‘Had it been possible to take up the economic plants of India by themselves, and to present a history of them to the English reader, we should at once have devoted ourselves to the task, with the certainty of obtaining an amount of encourage- ment which a so-called paying work is sure to command, but which one ofa more scientific nature is not thought worthy of receiving. We should, however, only be deceiving the public, were we to propose a scheme which, in the present deplorably backward state of scientific Indian botany on the one hand, and the confusion of Indian economic botany on the other, is literally impracticable: the difficulties have increased fourfold, from scientific botany not having advanced pari passu with the economic branch; and so long as plants themselves remain undescribed, it is obviously impossible to recognize what are useful, or so to define them that they shall be known by characters that con- trast with those of the useless. Our principal aim, however, being purely botanical, the most insig- nificant and useless weed is as much the object of our attention as the Teak, Sal, and tea: in the vegetable kingdom, and in the great scheme of nature, all have equal claims on our notice, and no one can predicate of any, its uselessness in an economic point of view.” “Every one who has studied Indian plants, whether for economic purposes or for those of abstract science, must have felt the want of a gener- al work which should include the labours of all Indian botanists, to be a very serious inconvenience. Our own experience in India has convinced us of this; for we found it often impossible to determine the names of many of the most ordinary, and, in an economic point of view, often most valuable forms; and every day’s additional experience in the preparation of this volume has served to show more and more clearly, that whilst such a work is wanting satisfactory progress is impossible. At present the student has to search in general system- atic works, for the descriptions of species; and as all of these are imperfect, a multitude of scattered papers must be consulted for the additions which have from time to time been made. These too have unfortunately so often been published without reference to preceding works of a similar nature, that the same plant has been described as new by xX FLORA MALESIANA [ser. I, vol. 4! many successive botanists, ignorant or neglectful of the labours of their predecessors.” So far HOOKER. To emphasize our inability to foresee practical results of taxonomic work I intend to mention a few recent instances in Malaysia showing that plants which seem useless at the present may stand in the focus of attention at a future date. ’ Twenty years ago it would have seemed the whim of a botanist to work on the species of a genus of foetid aroids, scientifically known as Amorphophallus. Few years later, however, the tubers of some species of this genus were found to be important commercially and industrially. The basic work on the distinction of the species, the notes on their distribution, their habit and structure proved to be most useful for agricultural purposes. The same holds for a genus of leguminous plants, Derris. The roots were found to contain a very valuable resin-like substance, rotenon, poison to fish and numerous insects but harmless to larger animals, also to man. As soon as its commercial value was recognized a sudden large demand for Derris rose. It soon appeared that not every species was valuable and so the original studies of Derris offered hold for a first segregation of promising material whereas the systematist was questioned about the characters by which the species could be recognized. The absence of any reliable taxonomic infor- mation of the genus Metroxylon prevents at present well-founded research on the economic possi- bilities of the sago-producing species which supply a basic food to the whole population of East Malaysia and Melanesia. Invariably it is the duty of the taxonomic bota- nist to supply basic data to research in directed (= applied) botany. In all cases the name of the species, and eventu- ally its varieties, is the alpha of knowledge, as it represents the key to existing literature embodying earlier work on habits, life-history, on distribution geographical and altitudinal, ecology and growth habit, current native names if any, efc. and Flora Malesiana must serve for this purpose. In the past e.g. tropical plant-breeding in some cases followed a wrong direction and might have achieved better results more rapidly when the aid of taxonomists had been available or requested. From the discussion of some selected topics above it will be clear that the taxonomic botanist in composing the Flora Malesiana will be able to offer critical knowledge of numerous forest prod- ucts, plants containing vegetable oils, fats, and resins, rattan, timber, gums, fruits, spices, insec- ticides, fibres, dyes, and medicines, or species which may serve for afforestation, for ornamental use, as new green manures, fodder plants, or possibly, species withstanding drought or being resistant to fire or inundation, suitable for combating erosion, and other economic aspects. In addition to taxonomical information, the Flora Malesiana will contain ecological data. In anthropogenic areas and eroded lands biological control of necessity will seek guidance in its com- prehensive survey of facts. Large amounts of money and energy have been wasted in the absence of professional planning, through negligence of funda- mentals. I remember attempts, as expensive as they were fruitless, of planting mangroves to protect the coastal area of a tropical harbour, a waste which would have been avoided when the ecological potentialities of mangrove forest had been duly considered. ! In (re-)afforestation, the choice of trees has to rely partly on previous experience, but directions can be given by field-taxonomists and by means of general rules of tolerance capacities. Native trees occupy in our forest-types fitting ecological niches, but it should not be assumed that they grow always under optimal conditions. An example is probably found in swamp forest trees which have roots tolerant of a very low aeration of the soil, a virtue not practically utilized, as far as I know, when planting on very poorly aerated soils. The ecological misunderstanding that all plants grow in nature under optimal conditions for their growth led to ‘forest plantations’ of quinine by JUNGHUHN. The Cinchona-crop was saved thanks to TEYSMANN who maintained that the plant should be grown in the open. Much trouble and still much more money could have been saved if this ecolo- gical principle had been better known. The Flora Malesiana is, therefore, of first in- terest to practice and may direct new research: it must give data as to where the plant occurs, in what quantity, under what life-conditions, and with what life-cycle. It ought to contain ecological and biological data, and a critical extract of the notes made by the collectors. None of us can pre- dict the industrial future of a neglected plant spe- cies, but we should be prepared for any coming rush on the botanical wealth of this vast archi- pelago, linking the Asiatic and Australian con- tinents. The aim of the Flora Malesiana is to compile a critical knowledge and a botanical standardization of the Malaysian flora of basic importance both to pure and to economic botany. How much of the flora is known? Often it is assumed—the majority of botanists being ac- quainted with the state of knowledge in Europe or North America—that the flora of these islands is sufficiently known, and the actual facts cause astonishment. For instance, not even the number of species is known otherwise than by very approximate cal- culation; 25.000 to 30.000 species of flowering plants is a conservative estimate. The Orchidaceae alone claim about 5000 species. Java possesses more than 500 species of ferns. The number of different species of trees in Malaysia is about 3000. The total number of genera is near 2400. The largest genera are found among the Orchids, Dendro- bium with ca 1110 and Bulbophyllum with about 933 recognized species. This is indeed astonishing if compared with the flora of Holland where the whole native flora (1) Kustaanwas en mangrove (Natuurwet. Tijdschr. Ned. Ind. 101 (1941) 82-85). Dec. 1948] Introduction XI amounts to little more than 1000 flowering plants. Counting all trustworthy and up to date revi- sions together, about 5000 out of a total of 25.000- 30.000 species are now more or less critically known. It appears that the bulk of the work re- mains still to be done. The area covered by the Flora Malesiana will besides Indonesia also include the Malay Pen- insula, Sarawak, Brunei & British North Borneo, the Philippines, Christmas Island, Portuguese Timor, and the whole of New Guinea (fig. 1). aS" = — Fig. 1. Delimitation and main divisions of the flora of Malaysia. It may be asked whether this is not an unneces- sary extension of the task to include foreign border countries. To explain this it ought to be realized that the demarcation lines of natural units seldom coincide with political boundaries. As much as possible, however, the demarcation of a Flora should be based on scientific, that is, plant geographical limits. Plant geographically the natural demarcation lines of the Malaysian flora pass through the isth- mus of Kra, between the Philippines and Formosa, and through Torres Straits, and include the Louisi- ades and the Bismarck Archipelago. An extensive geographical survey of the distribution of the Malaysian flora will be published in the 3rd volume of this work. The outcome! is wholly in confir- mation with the suggested demarcation lines which were drawn first, as I have mentioned, about a century ago by ZOLLINGER.? In the NW quite a number of typical Malaysian genera of forest plants fail to occur any further in the Indochinese Peninsula, e.g. Rafflesia, Rhizan- thes, camphorwood (Dryobalanops), benzoin (Sty- rax benzoin), kauri or copal (Agathis), true ironwood (Eusideroxylon), menggaris (Koompassia), etc. The Philippines possess an essentially Malaysian flora, in contrast to Formosa’s Japano-Chinese (1) Tijdschr. Kon. Ned. Aardr. Gen. 65 (1948) 193-207, 7 fig. (2) Natuurk. Tijdschr. Ned. Ind. 13 (1857) 293-322. floral character which was definitely demonstrated by MErRRILL.? The flora of New Guinea was formerly assumed to be essentially Australian in character. This in- terpretation was mostly based on zoological argu- ments and on the occurrence of few but very striking examples of plants which later appeared to be also spread westwards in the Moluccas and Celebes. O. WARBURG, in 1891,4 on account of important statistics, already showed the essential Malaysian character of the Papuan flora. Technically the botanist must in each case —whether the Flora Malesiana is limited to a political or to a natural demarcation—study and compare critically all species of the natural phyto- geographical unit. Plants described hitherto only from East New Guinea almost certainly occur also in West New Guinea, numerous species originally described from the Philippines occur in Celebes, the Moluccas and New Guinea, and the same holds for the Malay Peninsula, where the flora is inti- mately allied to that of Sumatra and Borneo. In identifying plants of Malaysia in the narrow sense, that is, limited to the Netherlands Indian boun- daries. the botanist is always obliged to revise or critically to take into consideration the species described from the border areas. This will cost him about the same time and labour as when admitting them into the final work. If these species are omitted, the Flora Malesiana will doubtless be out of date early and unneces- sarily. Bibliographic advantage of the Flora Malesiana. The absence of any definitely indicated centre of publication for Malaysian plants has led to a rather chaotic taxonomic literature. At the present moment revisions of Malaysian plants are pub- lished more or less frequently in about 10 impor- tant periodicals scattered all over the world, and occasional publications are found in some 50 others. An annotated list of former revisions will be presented in this volume to facilitate future study. No single individual can be supposed to own these journals and it is thus more or less private knowledge to those, who have access to a well-stocked library. In Malaysia there are only two libraries where they are nearly all represented, viz at Buitenzorg and Singapore. This is of course a rather unsatisfactory situation to naturalists, foresters, agriculturists, phytoche- mists, veterinarians, pharmacologists, and inter- ested private persons desirous tostudy the flora according to the best available data. The Flora Malesiana will put students of systematic botany generally in possession of the essence of literature. Sequence of publication. It is commonly under- stood that in a flora the sequence of publication ought to be in agreement with the ‘natural system’. This has been—I feel sure—a serious obstacle mentally and practically to all those who, pre- viously, have considered the project of this flora. Arguments against this sequence are in the first (3) Bot. Jahrb. 58 (1923) 599-604. (4) Bot. Jahrb. 13 (1891) 230-455. xII FLORA MALESIANA [ser. I, vol. 41, Dec. 1948] place the existence of several ‘natural systems’; it is tacitly agreed that the last word in ‘the natural system’ will probably never be spoken. A system now adopted may be obsolete when this flora is finished. A choice seems, therefore, difficult, as most of the systems are advanced by leading botanists who among themselves, may claim little priority of preference. It would be possible that the editors of the Flora Malesiana advance a system of their own. However, this falls beyond the scope of this Flora which is solely intended as a practical work. This technical difficulty, which was already mentioned on p. viii, in connection with the adop- tion of a system is a serious obstacle to the progress of the work. Clearly not at every moment a specialist is available for every family of flowering plants. This is more or less a matter of chance. Rapid and regular publication is most desirable and so every opportunity should be made use of. A ‘natural system’ consequently involves the ‘waiting’ of some manuscripts for many years because it is not yet their turn to be printed, and several volumes will be set up in one part but can be continued only at a remote period because for the ‘following’ family no specialist was available. The real disadvantage can be observed in works like the Flora of North America, in course of publication, of which, in 1941, were published 2 complete volumes and 55 loose parts belonging to 17 of the remaining 32 planned volumes. The same has been the case with the Flore Générale de I’Indo-Chine where most volumes ranged over a period of about 30 years before they were completed and could be bound. In the meantime consultation was very difficult because the indexes appeared naturally in the final instalment. The handling of the loose parts is undesirable both from a bibliographical and a practical standpoint. In the newly started Flore de Madagascar the families are numbered according to the natural system and are separately published and paged. The idea is that after completion the subscribers can arrange them into sequence and bind them accordingly. We must be aware, however, that this will hardly bring any advantage as the number of families in the Malaysian flora is 211, and that among them 70 families are represented by less than about 10 species, so that also in this case one has to handle a large amount of small unbound fascicles. A long time is needed to complete the Flora Malesiana, about 25 years at least.! This is cer- tainly not overestimated if compared with floras of similar magnitude as Flora Brasiliensis (1840- 1906), Flora of Tropical Africa (1868—hodie), Flora Capensis (1894-1933), Flora of British India (1855- 1897), Flore générale de Il’ Indo-Chine (1907—-hodie), Flora Australiensis (1863-1878). The exact duration cannot be calculated, this (1) Under the most favourable conditions as regards funds, and co-operation. depends largely on opportunity and facilities, and the joining of forces. The editors are fortunate in having received the promise of much co-operation, and they hope to be able to extend their resources still more. Moreover a considerable amount of recent publications exists which may easily be adapted to the flora. The here adopted scheme of ‘opportunity se- quence’ in the production of family revisions will remove any delay caused by the ‘natural system’. The addition of an up to date index to the contents of prior parts on the cover of each new appearing instalment will serve to verify in a moment if a desired group has already been revised. The size of the families is of course widely differ- ent ranging from 1—5000 species. At least one figure illustrating characteristics will be added to each family and large genus. The volumes will not exceed 500-600 printed pages. They must be easy in the hand, agreeable to work with, and bound in covers which may not be attacked by tropical insects, as we hope that numerous subscribers will be found in the Old World tropics outside the official institutions. Completeness of the Flora. No perfection can ever be attained in any tropical flora. Always novelties and new localities will have to be recorded. No squadron of botanists can ever comb a tropical area engirdling !/7 of the equator. Although completeness is a first aim set for this work, its future value will depend mainly on the amount of critical original study which it contains. The Floras of British India? and Australia are now definitely incomplete, but they remain first class sources of information. BACKER’s Schoolflora voor Java, of 1911, still meets present demands nearly as well as at the time of its appearance. If we can keep our flora to so high a standard it will become the keystone to future Malaysian systematic botany. The Flora Malesiana will be started with the flowering plants (Series J). Series II will comprise the ferns and fern allies and is estimated to occupy 3 volumes. Series ITI will be devoted to mosses and hepatics. These will take about 5 volumes. Series IV will treat the fungi and lichens. The number of volumes can as yet not be estimated. Series V is intended for the algae and other groups of unicellular cryptogams. For the series II-V special editors will be ap- pointed. The general method of treatment may possibly deviate somewhat from the first and largest series but the needs of these can hardly be estimated at the moment. Cc. G. G. J. VAN SZEBNIS Buitenzorg / The Hague, Sept. °44/ July °47. (2) Dr K. Biswas calculated that to the ‘Flora of British India’ consisting of ca 14000 species, ca 2000 have been added since its publication, a sur- prisingly low number in relation to its vast surface and variety of vegetation types (Proc. 30th I.S.C. pt II, sect. V, Bot., Pres. addr. p. 109). GENERAL*CONSIDERATIONS We should endeavour to determine how few, not how many species are comprised in the Malaysian flora. In writing the following chapters I have kept in mind the exemplary ‘Introductory Essay’ of J. D. Hooker in his ‘Flora Indica’ (1855), the precursor of the ‘Flora of British India’. For the same reasons that moved Hooker, I felt obliged to introduce the Flora Malesiana prop- er by some general considerations especially in- tended for co-operators less fortunate than I have been in acquiring an experience of long standing in the field. I may add that field experience often is invaluable when studying dried, always frag- mentary, materials in the Herbarium. Some of the subjects HOOKER treated are now too large to be included in one essay and, there- fore, the survey of the Malaysian collections, the physiognomy of the vegetation, and the genetic and floristic plant geography occupy the (introdtictory) volumes 1-3 of this work. The present essay will be entirely devoted to topics directly bearing on the study of systematic botany. Some of them I have previously discussed, or touched on, in my study of the origin of the Malaysian mountain flora.! As my intention is to further the study of Malaysian botany, I shall discuss only points of which a clear understanding is essential to the Malaysian naturalist. I will try to illustrate each case by reference to plants of this region. These points are: individual variation and racial segregation, variation caused by the environment, the problem of speciation and specific centres, hybri- dization, views on the status of the species and subspecies, migration and adaptation, and the way to interpret these concepts. These theoretical points are inseparable from a philosophical study of plants, and I believe it to be essential for systematists to explain the prin- ciples which have guided them in the execution and design of their work. HOoKER’s general instructions have guided me in my work, and I am convinced that in the flux of botanical conceptions in general aspects the words of the Master still hold their own. I desire to express here my admiration for this classic work by quoting /iteratim some passages of his essay. “Tt may seem almost chimerical to look forward to a time when all the species of the vegetable world shall have been classified upon philosophical princi- ples, and accurately defined; and it must be con- fessed that the present state of descriptive botany does not hold out much prospect of the realization of so very desirable an object. This, we think, is in a great measure due, not to any want of students willing and anxious to take up the subject, but rather to a gradually increasing misapprehension of the true aim and paramount importance of systematic botany, and of the proper mode of pur- suing the study of the laws that govern the af- finities of plants. We are therefore desirous, at the outset of a work which is devoted to these subjects, (1) Bull. Jard. Bot. Btzg III, 13 (1935) 358-407. of explaining our views on them; and as we trust that our work will fall into the hands of many beginners who are anxious to devote themselves usefully to the furtherance of botanical science, but who have not an opportunity of acquiring in any other way its fundamental principles, we shall make no excuse for dwelling at some length on the subject. We are also anxious to refute the too com- mon opinion (which has been productive of much injury to the progress of botany) that the study of systems presents no difficulties, and that descriptive botany may be undertaken by any one who has ac- quired a tolerable familiarity with the use of terms.” “‘There can be no doubt that any observant per- son may readily acquire such a knowledge of ex- ternal characters, as will in a short time enable him to refer a considerable number of plants to their natural orders; though even for this first step more knowledge of principles is required, than to make an equal advance in the animal kingdom: but to go beyond this,—to develop the principles of classification, to refer new and obscure forms to their proper places in the system, to define natural groups and even species on philosophical grounds, and to express their relations by characters of real value and with a proper degree of precision, de- mands a knowledge of morphology and anatomy and often of physiology, which must be completely at command, so as to be brought to bear, when necessary, upon each individual organ of every species in the group under consideration. To follow the laws that regulate the growth of all parts of the plant, especially the structure of stems, the func- tions of leaves, the development and arrest of floral organs, and the form, position, and minute ana- tomy of the pollen and ovule, and to trace the whole progress of the ovule and its integuments to their perfect state in the seed, ought all to be fa- miliar processes to the systematic botanist who proceeds upon safe principles; but no progress can be made by him who confines his attention chiefly to the modification of these organs in indi- vidual or natural orders.’’—So far HOOKER. Variability in characters of minor importance and description of extreme forms have led to a rather confused state of affairs. I believe that among the scores of species described many microspecies should be reduced to a much smaller number of true species, with a normal area of distribution and a normal variability in characters typical for Lin- nean populations which are intermediate between the species of extreme ‘splitters’ and extreme ‘lum- pers’. Much ‘splitting’ has been caused by de- scribing single extreme forms not exactly agreeing with the type or type-description; for practical purposes it is sometimes required to describe such forms as new species and to recognize them pro- visionally as new ‘entities’; the author’s conscience and eagerness to finish his task are thus tempo- rarily satisfied. This method has proved a failure and a serious handicap to the progress of tropical plant knowledge. XIV FLORA MALESIANA [ser. I, vol. 4! There are three methods of handling new col- lections, all being equally unsatisfactory. Firstly, provisional rapid identification of the material as to genus, or to species as far as is possible, and its insertion in the herbarium; collectors in general do not favour this method as only few final names can be provided onacursory examination. Secondly, a collection may be worked through by rough com- parisons to named specimens and with standard literature. This second method is rapid but all ex- treme forms and forms belonging to large genera or to difficult families which cannot be identified rom the available literature, are described as new (specimen description). By this method collectors get immediate results but science is burdened with a host of ‘endemic’ species which, as experience has shown, disappear by the score when a thorough monograph is made. Thirdly, a collection may be thoroughly studied, delaying results, as the iden- tification of extremes means in nearly every genus a preliminary revision. Hooker continues (/.c.):—‘‘A knowledge of the relative importance of characters can only be ac- quired by long study; and without a due appreciation of their value, no natural group can be defined. Hence many of the new genera which are daily added to our lists rest upon trivial characters, and have no equality with those already in existence. A pro- neness to imitation leads to a gradual increase in their numbers, without a corresponding increase of sectional groups. Indeed, even when the sectional groups are well defined, and the genera in them- selves natural, a too great increase in the number of genera is detrimental, by keeping out of view those higher divisions which are of greater impor- tance. The modern system of elevating every minor group, however trifling the characters by which it is distinguished, to the rank of a genus, evinces, we think, a want of appreciation of the true value of classification. The genus is the group which, in consequence of our system of nomenclature, is kept most prominently before the mind, and which has therefore most importance attached to it.’’! (1) ““We may make our meaning more clear by a few examples. The genus Ficus is surely more natu- ‘ral than the subgenera Pogonotrophe, Covellia, Urostigma, &c, into which it has been subdivided. So with the genera Anemone, Hedyotis, Erica, Andromeda, and others which have been split into many by modern systematists.”” R. BROwN, G. BENTHAM, J. D. HOOKER and others, in all their works, laboured to keep this important principle in view, and to impress it upon others; they have, however, failed to check the prevalent tendency to the multiplication of genera. I add here other examples of genera occurring in Malaysia which are separated by trifling charac- ters: Voandzeia differs from Vigna only in fruit biology, viz its globular pods ripening subterrane- ously. In Urena and Pavonia now only one fruit character remains the decisive distinction, Dillenia and Wormia are distinguished only in their fruit biology, Berberis and Mahonia are distinct solely in the foliage, Kibessia and Pternandra differ only “The rashness of some botanists is productive of still more detrimental effects to the science in the case of species; for though a beginner may pause before venturing to institute a genus, it rarely enters into his head to hesitate before proposing a new species. Hence the difficulty of determining synonymy is now the greatest obstacle to the pro- gress of systematic botany; and this incubus un- fortunately increases from day to day, threatening at no very distant period so to encumber the science,” that a violent effort will be necessary on the part of those who have its interests at heart, to relieve it of a load which materially retards its advancement. The number of species described is now So very great, and the descriptions are scattered through such a multitude of books, that even after long research it is difficult to avoid overlooking much that is already known; and when botanists with limited libraries and herbaria institute new species, it is almost certain that the latter will be found to have been already characterized. To such an extent is this carried, that we could indicate several works, in which one half and even more of the species are proposed in ignorance of the labours of other botanists. Indian Botany unfortunately, far from forming an honourable exception in this particular, presents a perfect chaos of new names for well-known plants, and inaccurate or incom- plete descriptions of new ones’’. “It must be remembered too that the Linnean canon, by which twelve words were allowed for a specific character, is now becoming quite inade- quate to the requirements of the science; and that the brief descriptions, which are now so generally substituted for definitions, unless prepared with the greatest skill, as well as care, and after an inspection of very numerous specimens, seldom express accurately the essential characters of a plant. Itis indeed becoming more and more evident, that in the great majority of instances no definition is sufficient to enable inexperienced botanists to determine with accuracy the species of a plant, even when the whole genus is well known; much more is this the case in genera, many of whose species are yet undiscovered; and most of all, in those where the forms, though sufficiently well known, are liable to much variation. In the last case their determination becomes a special study; in unimportant characters of the calyx tube, efc. The separating characters are far less important than those which, in other genera, serve to divide sections or subgenera. The more species are described the more differ- ences originally accepted as of generic rank tend to disappear. A redefinition of the generic charac- ters is often delayed, and the attitude in ‘local- monographs’ is mostly to keep at all costs the old delimitation in order to avoid laborious mono- graphic work. Suggestive casual remarks are often made in local works, but decisions deferred. (2) In entomology this has led to an intolerable chaos (cf. The New Systematics 1940, p. 475-491). The same holds for several large groups of the Fungi. Dec. 1948] General considerations XV and when attempted without access to authentic specimens, leads to inextricable confusion, and its evil effects are not confined to specific botany, but extend to all departments.” “The pages of our Indian Flora will supply numerous illustrations of these remarks, and we would direct the attention of those commencing the study to the lesson to be derived from these instructive errors; for where the first botanists of the day have failed, beginners cannot be expected to succeed. It cannot be too strongly impressed upon all students of botany, that it is only after much preliminary study, and with the aids of a complete library, and an herbarium containing authentic specimens of a very large proportion of known species, that descriptive botany can be effectively carried out; and it would be well for science if this were fully understood and acted upon.” “The prevailing tendency on the part of students of all branches of natural history, to exaggerate the number of species, and to separate accidental forms by trifling characters, is, we think, clearly traceable to the want of early training in accurate obser- vation, and of proper instruction in the objects and aim of natural science. Students are not taught to systematize on broad grounds and sound principles, though this is one of the most difficult processes, requiring great judgement and caution; or, what is worse, they are led by the example if not by the precepts of their teachers, to regard generic and specific distinctions as things of little importance, to be fixed by arbitrary characters, or according to accidental circumstances. As a consequence, the study of systematic botany is gradually taking a lower and lower place in our schools; and, being abandoned by many of those who are best qualified to do it justice, it falls into the hands of a class of naturalists, whose ideas seldom rise above species, and who, by what has well been called hair-splitting, tend to bring the study of these into disrepute.” “We therefore earnestly recommend to the Indian botanist the detailed study of individuals and their organs with the view of determining their limits of variation.” WiGut and Arnott! formulated their warning to beginners as follows :— ““We shall perhaps be severely censured for cut- ting down species. We have all along considered it as trifling with nature to separate species on slight or variable grounds, nor could we ever under- stand the ‘cui bono’ for which so much ingenuity in splitting hairs has been wasted. Before we deter- mined what was a species, we examined with care numerous specimens from the same and different localities; and so far we have had an advantage over many other of the European botanists who have described Indian plants, they having only seen one or two isolated specimens. Numerous obser- vations too were made in the plants in their natural situation, the result of which went to prove, what we have frequently endeavoured to enforce by (1) Prod. Fl. Pen. Ind. Or. 1 (1834) p. xxxi. examplés throughout the present volume, that no precise shape of leaf or quantity of pubescence is of any value, although both of these seem in each species to be limited within certain variations. With regard to varieties, we have seldom distinguished any unless well marked and tolerably constant; we are aware, indeed, that these correspond to what some naturalists call species, but our own obser- vations have convinced us, that varieties and forms, as well as species, may be constant in similar situ- ations, and even in widely different situations, for many years, if raised from seeds either obtained from the original locality or from cultivated plants; the cultivated cerealia and garden vegetables ought to lead to such an hypothesis without any addition- al proof.’’ So far WiGHT and ARNOTT. HOOKER continues :— “In relative size especially, the observer will find immense variation; for, unlike the animal creation, proportional dimensions are of small moment in the vegetable kingdom. This fact, so familiar to the botanist of experience, is always a puzzle to the zoologist, who fancies he perceives a vagueness and want of exactness in all botanical writings (except in those of the too numerous class that make a parade of measuring to lines organs that vary inches), that contrasts unfavourably with des- criptive zoology. Symmetry is again only a relative term amongst plants, for even such leaves as grow in pairs are never alike, and often differ much in form, texture, and colour; whilst the various sepals, petals, etc. of an individual flower, never so exactly correspond as the relative members of an animal do; and there are still greater differences between these organs, when taken from different flowers.” “Tt is hardly necessary to allude to the desira- bility of studying the various forms induced by artificial causes: the browsing of cattle on shrubs, for instance, which is almost invariably followed, by an abnormal state of foliage on the subsequently developed shoots, has been a prolific source of bad species; while there is scarcely an operation of man that does not tend to produce change in the vege- tation surrounding him.” “It will generally be found that botanists who confine their attention to the vegetation of a cir- cumscribed area, take a much more contracted view of the limits of species, than those who extend their investigations over the whole surface of the globe. This is partly, no doubt, owing to the force of bad example; and partly to the fact that the student who takes up the study of the flora of his native country, finds that the species are all toler- ably well known, and that no novelty is to be discovered. There is therefore a natural tendency to make use of trifling differences, from the scope which they afford for minute observation and critic- al disquisition; whilst the more close comparison of the few species which come under his investi- gation, leads the local botanist to attach undue importance to differences which the experienced observer knows may be safely attributed to local circumstances. To this tendency there can be no limit, when the philosophy of system is not under- stood; the distinctions which appeared trifling to XVI FLORA MALESIANA [ser. I, vol. 4! botanists a quarter of a century ago, are at the present day so magnified by this class of observers, that they constantly discover novelties in regions which have been thoroughly well explored; con- sidering as such, forms with which our predecessors were well acquainted, and which they rightly re- garded as varieties. ! “Another result of the depreciated state of systematic botany is, that intelligent students, being repelled by the puerilities which they everywhere encounter, and which impede their progress, turn their attention to physiology before they have ac- quired even the rudiments of classification, or an elementary practical acquaintance with the charac- ters of the natural orders of plants. Unfortunately, in botany, as in every other branch of natural science, no progress can be made in the study of the vital phenomena except the observer have a previous accurate acquaintance with the various modifications under which the individual organs of plants appear in the different natural orders, and such an appreciation of the comparative value, structural and morphological of these modi- fications, as can only be obtained by a careful study of the affinities of their genera and species. Igno- rance of these general laws leads to misinterpre- tation of the phenomena investigated by the phy- siologist, and to that confusion of ideas which is so conspicuous in the writing of some of the astute physiological observers of the day.” “The modern system of botanical instruction attempts far too much in a very limited space of time, and sends the student forth so insufficiently grounded in any branch of the science, that he is unprepared for the difficulties which he encounters, let his desire to progress be ever so great. The history of botanical discovery, and the philosophy of its advance, form instructive chapters for the student in any department of natural science.” ““We owe to LINNAEUS the establishment of the doctrine of the sexuality of plants; and we find by the writings of the same great naturalist, that be- sides foreseeing many physiological discoveries, he preceded GOETHE in the discovery of morpho- logy, a doctrine which, more than any other, has tended to advance scientific botany. A third great discovery, that of the nature of the ovule, and the relation of the pollentube to the ovary, received its principal illustration at the hands of BROowN, our chief English systematist, and of BRONGNIART, also a practised botanist.’’ “It should not be forgotten, that the relative importance of physiology is very different in the animal and vegetable kingdoms. In the former, structure and function operate so directly upon one another, that the great groups are, to a certain extent, defined by well-marked external characters, which are at once recognizable by the student, and are familiar, or at least intelligible, to those even (1) “Many of the species which have been revived in modern times were indicated by HALLER, RAy, TOURNEFORT, and other ancient botanists, but were reduced to the rank of varieties, when the science was reformed by LINNAEUS.” who have paid no attention to natural history. In the vegetable kingdom this is by no means the case: the processes of assimilation and secretion present but little of that complication which ren- ders the study of animal physiology so important; they are, on the contrary, uniform almost through- out its whole extent, and moreover so simple in their modus operandi, that this very simplicity pre- vents their being rightly understood. In conse- quence, even the two great classes of Monocotyle- dons and Dicotyledons are not distinguishable without considerable practice and study; and were we dependent upon actual inspection of the organs whence the essential characters of these groups are drawn, for the means of recognizing, Systematic Botany would be an impracticable study.” “Herein lies one great obstacle which meets the beginner on the very threshold of his botanical studies: he sees the great divisions of the animal kingdom to be recognizable by mere inspection, and that familiar characters are also natural, and available for purposes of classification: the very names of the groups convey definite information, and to a great extent give exact ideas. Birds, fishes, reptiles, etc. are all as natural as they are popular divisions; but what have we in the vegetable king- dom to guide the student through the two hundred and fifty natural orders of flowering-plants? As with a new language, he must begin from the very beginning, and also avail himself of artificial means to procure as much superficial knowledge of struc- ture and affinity as shall enable him to see that there is a way through the maze. Hence the obvious necessity of an artificial system of some sort to the beginner, who has, at the same time, to master a terminology, which, if not so complex as that of zoology, is more difficult at the outset, from the want of standards of comparison between the or- gans of plants and those he is familiar with in himself as a member of the sister kingdom. Ap- plying these remarks to practice, the botanical student finds that he has much to unlearn at the very outset; in many cases he has misapplied the terms root, stem, leaf, etc., and contracted most erroneous ideas of their structure and functions; while he is startled to find that the popular di- visions of plants into trees, shrubs, and herbs, —leafy and leafless, water and land, erect, climb- ing, or creeping,—are valueless even as guides to the elements of the science.” ‘It is not however to be supposed, because pure physiology is of secondary importance to the right understanding of the affinities of plants, that botany is therefore a less noble or philosophical study than zoology; since we find anatomy, development, and morphology, occupying a very far higher rank in proportion. Being deprived, as he is in most cases, of all technical aids to the determination even of the commoner exotic natural families, the systematist is compelled to commence with the knife and microscope, and can never relinquish these implements. Systematic Botany is indeed based upon development; and no one can peruse, however carelessly, the most terse diagnosis of a natural order or genus of plants, without being Dec. 1948] General considerations XVII struck with the variety and extent of knowledge embodied as essential to its definition and recogni- tion. Not only are the situation and form, division or multiplication, relative arrest or growth, of the individual organs exactly defined, in strictly scien- tific and scrupulously accurate language, but the development of each is recorded from an early stage: the vernation and stipulation of the leaves; the aestivation of the young calyx and corolla, and their duration relatively to other organs; the de- velopment and cohesion of the stamens; the po- sition and insertion of the anther; its pollen; the cohesion or separation of the carpels, and the stages of their development from the bud to the mature fruit, and from the ovule to the ripe seed, are all essential points; all however minute, must in many cases be actually inspected before the position of a doubtful genus can be ascertained in the Natural System; and this is not the exception, but the rule.” “The necessity for acquiring so extensive and detailed a knowledge indicates a power of variation in those organs from which the natural characters are drawn, that defeats any attempt to render one, or a few of them only, available for the purposes of classification; and hence it is that the study of morphology or the homologies of the organs, be- comes indispensable to the systematist; by this he reduces all anomalies to a common type, tests the value of characters, and develops new affinities. The number, form, and relative positions of organs may supply technical characters, by which ob- servers of experience recognize those natural orders under which a great number of plants arrange themselves; but a knowledge of structure and ana- tomy alone enable the botanist to progress beyond this, and to define rigidly: whilst the study of development affords him safe principles upon which to systematize and detect affinities, and mor- phology supplies the means of testing the value of the results, and reveals the harmony that reigns throughout the whole vegetable world.” “Physiology, again, is a branch of botany very much apart from these: its aim is the noblest of all, being the elucidation of the laws that regulate the vital functions of plants. The botanical student of the present day, however, is too often taught to think that getting up the obscure and disputed speculative details of physiology, is the most useful elementary information he can obtain during the short period that is given him to devote to botany; and that, if to this he adds the scrutiny of a few of the points under a microscope, he has made real progress as an observer. This, we maintain, is no more botany, than performing chemical experi- ments is chemistry, or star-gazing astronomy. A sound elementary knowledge of vegetable physi- ology is essential to the naturalist, and should indeed be a branch of general education, as it requires nothing but fair powers of observation and an ordinary memory to acquire it. For the student to confine his attention to this knowledge of the vegetable world, and to try and improve upon it by crude experiments of his own, under- taken in ignorance of the branches of pure botany we have enumerated, is a very rational amusement, but nothing more.” “The students are indeed, in too many cases, perfectly ignorant of the elements of natural science, and require some practical acquaintance with plants and their organs, before they can ap- preciate the relations of the different branches of botany to one another, or discriminate between what is essential to understand first, and what is better acquired afterwards. Were the elements of science taught at schools, this would not be so: we should then have the student presenting himself at the botanical lectures fully prepared for the more difficult branches of science, and for making that progress in them for which the professor’s aid is indispensable. A sound practical knowledge of system we hold to be an essential preliminary to the study of the physiology of plants—a study which requires also a practical acquaintance with organic chemistry, consummate skill in handling the dissecting knife, and command over the micro- scope, a good eye, a steady hand, untiring per- severance, and above all, a discriminating judgment to check both eye, hand and instrument. A combi- nation of these rare qualities makes the accom- plished vegetable physiologist, and their indispensa- bility gives physiology its pre-eminence in practice.’’ “It has been with no desire of obtruding our views upon our readers that we have ventured to discuss these obscure subjects with relation to Indian plants, but from a conviction, that in the present unsatisfactory state of systematic botany it is the duty of each systematist to explain the principles upon which he proceeds; and we do it not so much with the intention of arguing the sub- ject, as of pointing out to students the many funda- mental questions it involves, and the means of elucidating them.” “To every one who looks at all beneath the surface of descriptive botany, it cannot but be evident that the word species must have a totally different signification in the opinion of different naturalists; but what that signification is, seldom appears except inferentially. After having devoted much labour in attempting to unravel the so-called species of some descriptive botanist, we have some- times been told that the author considers all species as arbitrary creations, that he has limited the forms he has called species by arbitrary characters, and that he considers it of no moment how many or how few he makes. So long as this opinion is founded on conviction, we can urge no reasonable objection against its adoption; but it is absolutely necessary that the principle should be avowed, and that those who think the contrary should not have to waste time in seeking for nature’s laws in the works of naturalists who seek to bind nature by arbitrary laws. So again with regard to specific centres; except we are agreed with an author as to whether the same species has been created in one or more localities, and at one or more times, we shall be at cross purposes when discussing points and principles relating to identity of species and geographical distribution.” “Great differences of opinion have from the XVIII FLORA MALESIANA [ser. I, vol. 4! earliest days of science always existed on the nature of species. The prevalent opinion has undoubtedly at all times been, that a species is a distinct creation, distinguishable from all others by certain perma- nent characters. Many eminent philosophers, how- ever, have taken a contrary view; of these the best known have been LAMARCK, and more recently the anonymous author of the ‘Vestiges of Creation’.” —So far HOOKER. Modern biological science has progressed rapidly in the last decades through the results of experi- mental genetics. Though it is far from easy to weld the often contradictory opinions into a satisfactory whole, views relating to matters of variation have much gained. In the following pages I will try to discuss on this new basis the value to be attributed to characters of less than specific importance and a number of considerations which may lead to increased accuracy in judging specific delimitation. Trifling characters, such as peloric and cleistoga- mous flowers, have led to the creation of worthless new genera; galls, insect bites, and parasitic fungi have been mis-interpreted and caused the publi- cation of new species of Phanerogams. Individual variations, either intrinsic or extrinsic have, in a similar way, induced systematists working on tropical plants to distinguish more species than Nature intended. It is not our intention to limit phytography to a merely administrative function in the study of botany, but to treat it as an essential of natural philosophy. The systematist ought to keep pace with cytogenetics, physiology and morphology, ! phytochemistry, phytogeography, ecology, genetics, i. e. experimental taxonomy. Inadequate material? and information are the chief causes which prevent the phyto-systematist from applying the results obtained by these branches of botanical science. The systematist is seldom favourably regarded by the layman or student of directed botany. They are opposed to changes in nomenclature, being unable to gauge the force of the arguments for a ‘new’ name for a familiar plant and so rarely accept the judgment of taxonomists. When, on the other hand taxonomic problems are tackled by applied scientists nomenclature and specific distinction become chaotic; entomology, mycology, forestry, agriculture and horticulture supply many examples. A wish for simplification, impatience, or even per- sonal vanity or the desire for pecuniary gain have caused hosts of ‘species’ to be added to our lists (1) For the value of wood anatomy in taxonomy, see DEN BERGER, in Handel. 4e N.I. Natuurwet. Congres (1926) 397. (2) Cf. WIGHT, in a letter to GriFFITH, dated April 15, 1842:—‘“‘How people can work on dry plants I cannot imagine. I am daily convinced of the poverty of the study from such materials, unless a man has seen much of living structure.” MIQUEL ignored this remark, and on sterile and inadequate material based a host of species from Sumatra which even at present are not wholly elucidated. by applied workers. Not long ago a forestry officer made a study of Agathis? in Malaysia in which 13 species and 2 doubtful ones, that is 15 entities, were distinguished. In the same material the late Dr Danser, whose judgment and experience cannot be doubted, distinguished only 3 divergent species with a number of local geographic variations. He found it very difficult to define the latter. Additional material showed that the keys and distinctions presented for the 13 species did not hold to the satisfaction of the Forest Research Station, from which this work emanated. In plant families of economic importance particularly in Gramineae, Rutaceae, and Leguminosae, similar work has re- sulted in multitudes of microspecies provided with binomials; by such a proceeding nothing is gained and much lost. An example of the difficulties arising between taxonomy and an applied science when a good revision is absent, is the following:—a Clausena of unknown origin was cultivated for economic purposes at Buitenzorg. I referred it to Clausena anisum-olens (BLCO)MERR. but the phytochemist was dissatisfied, the properties of the oil did not tally with data recorded from the same species in the Philippines. I then sent ample material with full notes to Dr TANAKA, Dr SwWINGLE, and to the Kew and Paris Herbaria, for identification. The answers were all different and the phytochemist was, of course, disgusted with the practical results of taxonomy, because now he had the choice among 5 names for his plant. By way of comfort I expressed the hope that a systematist would some day make a satisfactory monograph of the genus.+ In order not to raise his hopes too high I remarked that even then some research from him would be needed to establish the assumed constancy of the oil properties as a specific character. I also inform- ed him that taxonomy has sometimes scored by predicting phytochemical facts, e.g. when HALLIER supposed the presence of valerianic acid in Vibur- num> on phylogenetic grounds only. In the following two chapters general infor- mation on variation as a source of superfluous binomials is collected for the benefit of those with no field experience of the Malaysian flora. I distinguish variations induced by the environment from those belonging to the genetic composition of populations, and I have tried to illustrate them by examples in Malaysian phytography. Often the number of examples is too small, and chapters overlap, but in the course of time every student of Malaysian botany will meet with other equally telling cases. May they stimulate the wish to avoid lapses of this character by conscientious treatment of the revisions in Flora Malesiana. (3) Bull. Jard. Bot. Btzg III, 16 (1938) 455-474. (4) Compare R. WIGHT in a letter to GRIFFITH, dated March 30, 1841:—... ‘‘as you say Botany is difficult, and increasingly so, but Botanists are to blame for this. No remedy will be so effectual as the publication of Monographs.” (5) Med. Rijksherb. Leiden no 14 (1912) 36; ibid. 37 (1918) 92. Cf. also V. valerianoides ELM. Dec. 1948] General considerations XIX aR ATITONS MOSTLY INDUCED BY THE ENVIRONMENT (Phenotypic modifications) Phenotypic modification is the response to environ- mental conditions, such as climate, soil, exposure, altitude, temperature, wind, fire and living organ- isms. The genetic qualities govern the character of the plant, but the environment in which the plant develops determines the actual and final appear- ance of the individual. The changes or differences from the ‘normal plant’ are called modifications. Such changed characters are not themselves in- herited, however, though the manner in which a plant reacts to environmental conditions is. In some cases an external change may be reversed by a change in the environment during the develop- ment of the individual but in other cases, when factors act in the seedling stage only, the effects in the individual are irreversible. It is necessary to agree about the concept ‘normal plant’. This is far from easy, as each speci- men grows under a different combination of CEB- factors (Climatic, Edaphic, Biotic). We might ap- Intrinsic a 1 2 Ontogeno-morphosis . ji 5 6 7/ Teratologo-morphosis . Hypselo-morphosis Photo-morphosis . Climatic Hora-morphosis . 2s , Anemo-morphosis oy IS: Edapho-morphosis Hydro-morphosis . ney led Phyto-morphosis . Nees Zoo-morphosis . Ae 20: | == | 11 8) ( Bi | , Anthropo-morphosis . < 23. Ontogeno-morphosis 1. Juvenile forms Juvenile forms often differ widely from the mature plant. Seedlings of many Leguminosae differ greatly from the adult in foliage and other charac- ters. The youth form of Cassia javanica L. pos- sesses large metamorphosed twigs acting as thorns (1). Thorny juvenile specimens are also found in Alangium. In general, flowering twigs have smaller leaves than sterile branches; this often gives rise to diffi- . Juvenile forms : : : . Precocious flowering ( Pacdoeeaesy : F : : Xxi proach the idea by saying that ‘“‘the normal plant results froma genetically average individual under average natural environmental conditions”’’, average to be understood in the sense of optimal. This ‘normal’ individual is never a reality but remains an abstraction. Though the difference between phenotypic and genotypic variation is clear, the field botanist—and still more the herbarium botanist—is not always able to recognize it. Only experiments may furnish proof. For instance a dwarf shrub in an area sub- ject to fire or browsing animals may assume this stunted form through these CEB-factors but it is also possible that the stunted form is a specialized race adapted to these conditions and thus selected by nature itself from the specific population. Ex- perimental breeding must decide its constancy. I have arranged the phenotypic modifications under several headings—which partly overlap and interlock—in the following sequence: p. xix . Dimorphous foliage XXii . Dimorphous seeds and fruits XXIV . Dimorphous flowers XXV . Cleistogamous flowers XXV . Teratological forms XXV . Phenotypic effect of altitude XXVI . Epiphytes XXVili . Shade forms . XXIX Influence of drought XXIX Seasonal variation XXIX Wind forms XXX . Fumarole plants XXX . Rock plants; calcareous and silicious soils XXx1 . Solfatara plants XXXili Water and swamp plants XXXIV Fungus and bacterial diseases, and symbiosis XXXV . Ant plants (myrmeco-morphosis) XXXV Galls deceptive to phytographers ( cecidio: morphosis) 5 XxKvi 21. Influence of browsing animals (pascuo-morphosis) XXXVI 22. Influence of fire (pyro-morphosis ) . XXXVIi Pioneer plants ; . XXXVili . Savannah trees . XXXViil culty in identifying non-flowering material and s one of the pitfalls if new species are based on sterile material. An example is Campnosperma acutiauris BoERL. & KoorRD. (Anacardiaceae) described on sterile juvenile material from Sumatra. The leaves are large and conspicuously auriculate-amplexi- caulous. A similar juvenile form was later found in West Java, together with mature trees. These possessed much smaller non-auriculate leaves (fig. 2). The plant appeared to represent a species of Tristania (Myrtaceae)(2); its specific identity will probably remain obscure, however, as several species produce similar juvenile forms. XX FLORA MALESIANA [ser. I, vol. 4! Youth forms of Myrica longifolia T. & B. differ strongly from mature specimens in possessing distinct stipules and incised larger leaves. Incised leaves of seedlings occur in a score of arboreous plants, e.g. many Bignoniaceae, Pro- teaceae, Gmelina, Lonicera, Alangium, Vitex, &c. Leaves of young trees of Pangium edule REINW. Fig. 2. Large leaf of a juvenile specimen, small leaves of a mature tree of Tristania sp. (Myrt.) in W. Java, x 1/3. are 3-lobed whereas the leaves of mature trees are entire. The growth of different parts of the plant is often very disproportionate. In some Symplocos species I found the leaf teeth were mature and large in young leaves but inconspicuous in mature foliage: they possibly have some (?excretive) function during youth only. BACKER (3) found the leaf tip earlier developed than the blade in some species of Dioscorea; it disappears also sooner. A peculiar development occurs in the growing leaves of some Meliaceae, e.g. Chisocheton (fig. 3). Very peculiar juvenile forms greatly differing from the later normal foliage, have been describ- ed in various climbing plants such as some spp. of Adenia, Medinilla, Macrozanonia, Piper, Araceae, Ficus, &c. Juvenile specimens of these trunk climbers are always sterile. Their foliage is mostly broader than that of mature plants, and is appressed to rocks or tree trunks. The similarity in their appearance may cause considerable con- fusion as e.g. is shown by the type specimen of Ficus peltata BL. which was recently proved to represent a juvenile specimen of some climbing species of Piper. In several Malvaceae, Leguminosae, Sterculi- aceae, Tiliaceae, juvenile leaves are often different from mature ones (fig. 4). Sterculia polyphylla R. Br. is a juvenile stage of St. foetida L.; young trees often possess leaves having 10-15 narrow leaflets, mature trees have mostly 5—9-foliolate leaves with broader segments. Young Lasia spinosa THw. is very different from the mature plant. Ficus basi- dentula Mia. is merely the juvenile form of F. cal- losa WILLD.; it is quite common in the hedges at Buitenzorg. The polymorphy in the habit and foli- age of Ficus quercifolia Roxs. and F. heterophylla L.f. is unbelievable. In juvenile forms of Nepenthes the shape of the pitchers may considerably differ from that in mature plants; asa result juvenile Nepenthes cannot with certainty be identified. In Carallia lucida Roxs. leaves of mature trees are oblong to obovate with very shallowly serrate to entire margins; saplings, however, have oblong to lanceolate leaves distinctly serrate (4). Other cases of old mature plants differing from young ones are found among lianas in which the shape of the stem may change considerably: Cissus tuberculata Bu. has terete tuberculate stems but they later become flat and, in older stems, up to 60cm broad looking like gigantic ribbons! The latter were described as a separate species, Vitis lanceolaria WALL., but the two forms are merely two stages of one species. The stems of lianas generally change greatly with age, through the de- velopment of corky warts and wings, together with secondary wood not present in young flowering twigs. Spines sometimes disappear in lianas and trees with age; in some cases, on the other hand, they enlarge considerably. A peculiar case is repre- sented in two undescribed Cucurbitaceous lianas from the Lesser Sunda Islands, both having a Fig. 3. Growing leaf tip of Chisocheton sp. (Meliac.) (bbs 23227): ast Dec. 1948] General considerations XXxI swollen base; in Gynostemma sp. this ‘podagric’ base is smooth, in A/somitra sp. it is spiny (5). Habit also sometimes changes with age: Anci- strocladus is sometimes a shrub in youth whereas later it becomes distinctly scandent. Climbers which have no support may sometimes grow into semi-erect shrubs; this I once observed in a plant Fig. 4. Hibiscus sagittifolius KURZ (Malvac.), leaves from one specimen (Indramajoe, W. Java), x 1/2. of Smilax modesta DC. in a grass field on Mt Diéng. Spotted leaves often occur only or predomi- nantly in juvenile specimens. Strobilanthes picta KoorD. was a new species proposed on account of silvery spots on the leaves. However, it is a juvenile stage of Str. cernuus BL. Similar cases are known . in Begonia, Cissus, and other genera where these spots may disappear with age. In greenhouses these juvenile forms are preferred for ornamental purposes. ; Juvenile forms of plants with pinnate leaves sometimes have a much larger number of pinnae e.g.Campsis pandorana (ANDR.)STEEN. c.n. (6). An example of a new species based on a juvenile plant is found in Dacrydium: the type specimens of Dacrydium junghuhnianum Mig. from Sumatra consist of juvenile specimens of D. elatum WALL. with long loosely set needles. In the herbarium flowers sometimes open during drying and create the impression of being mature. This is specially the case with flowers having val- vate terete corollas e.g. Symplocos § Cordyloblaste, Styrax, Polyosma, Proteaceae, Loranthaceae, &c. MIQUEL described a new species of Lonicera from Sumatra L. sumatrana Mia. In his description short corollas are mentioned; owing to this mistake the species was subsequently placed in the wrong sec- tion and described twice again, once from Burma and once from Sumatra (7). The examination of MIQUEL’s type specimen revealed that he described immature flowers, in fact buds which had opened in the herbarium. Immature woody capsules or strobili of Myrtaceae, Theaceae, Coniferae, Casua- rina, &c. also tend to open after drying. There is often a great similarity in the leaves of watersprouts of mature trees with those of saplings: large size, deeper incised teeth, thinner texture, e.g. in Symplocos, Ficus, Sapotaceae, Dipterocarpa- ceae, etc. A still unsolved case is that of Evonymus japo- nicus THUNB. of which a sterile slender climbing and rooting form is frequently found in the Javan mountain forests. I originally took it for a juvenile shade form (8). Not until 1941 did I succeed in finding it flowering and fruiting on the open sum- mit of Mt Jang. It is unknown whether the shade conditions in the juvenile stage determine the later morphology. Cited literature: (1) A. J. KoENS, De Trop. Na- tuur 2 (1913) 174; see also Koorpers, Bull. Jard. Bot. Btzg. III, 1 (1919) 168. (2) Tectona 22 (1929) 1336-1340. (3) Handboek Flora Java pt 3 (1924) 109. (4) Schoolflora voor Java (1911) 486. (5) Fig- ured in De Trop. Natuur 29 (1940) 6. (6) Bignonia pandorana ANpbR. (7) Journ. Arnold Arbor. 27 (1946) 441, 445. (8) De Trop. Natuur 22 (1933) 175-176. 2. Precocious flowering (paedogenesis) In several Malaysian plants pre- cocious flowering is observed. Costerus (1) recorded flowering seedlings in Melia arguata DC. (fig. 5). BAcKER found them in Melia azedarach L. and J.J. SMITH described (2) the same pheno- menon in Murraya paniculata L. In Cocos nucifera L. precocious flowering is often seen. The late Dr A. RANT observed flowering seedlings in Cinnamomum zeyla- nicum THw. (oral comm.). Other plants in Malaysia in which pre- cocious flowering has been observ- ed are Swietenia mahogani JACQ., Coffearobusta, Citrus decumana L., Nicotiana tabacum L., Sesbania sericea DC., Vigna sinensis ENDL., Teramnus labialis SPRENG., Tectona grandis L.f., Kalanchoe pinnata Pers., and Ailanthus sp. In plants which flower strictly periodically precocious flowering is sometimes controlled by the Fig. 5. date of sowing. If sown too late Precocious they flower together with full- flowering grown plants sown earlier. This (paedogenesis) is a fact well-known to agricul- in Melia turists (in Java e.g. in Hibiscus arguta DC. Spp.). (Meliac.). Precocious flowering may also (after be caused by poor soil or some COSTERUS) XXII methods of pruning. An example of the former cause is Osbeckia pusilla ZOLL. which is a flowering dwarf of O. chinensis L. on poor soils. Sometimes dwarfed plants flower when very small and represent distinct varieties or strains, e.g. the dwarf of Canangium odoratum BAILL. f. pumila (3) grown in pots in Malaysia (introduced Z Wes. GED 2 GA yee ‘s Gees WIE = 5 we j oe ps a OF: } Le Fig. 6. Monophyllaea horsfieldii R.Br. (Gesn.), adult plant, one cotyl large and leafy, the other (in front) bract-like, soon disappearing (W. Java, Kalapa Noenggal), x 2/s. from China), Aglaia odorata Lour. var. microphyl- lina DC., and a dwarf of Punica granatum L. The skill of Chinese and Japanese horticulturists in raising dwarfs is due partly to the selection of pygmy varieties but more important is their skill in impoverishing the plant without starving it (4). Many dwarfs are found near solfatara, on rocks, and on silicious soils (cf. § 15-16). Flowering juvenile forms are comparable to the neoteny found in the animal kingdom. Pteridophytes generally are apparently more plastic with regard to precocity than Phanerogams, FLORA MALESIANA [ser. I, vol. 4! and several species are known to form spores in dwarf or juvenile specimens which have some- times been described as distinct species. It has been assumed that pygmy species in Antrophyum may possibly represent neotenous stages of other species. COPELAND described in 1939 (5) a dwarf fern from Borneo as Holttumia, but it is Dr DonxK’s conten- tion that this fern is a precocious stage of a Tae- nites. In the genera Teratophyllum, Stenochlaena and Lomariopsis, HOLTTUM (6) was able to demon- strate that a great deal of confusion is caused by the description of juvenile stages; being familiar with the living plants in the field he clarified the true status and affinities of a number of obscure species. DrELs (7) compiled an instructive book on juve- nile forms, giving instances where the juvenile foliage persists in the mature flowering plant, a course of development comparable to neotenous forms in zoology. Australian and New Zealand botanists have written a great deal about this phenomenon of heteroblasty which in those floras has apparently an important bearing on speciation (8). I cannot remember a Malaysian plant sus- pected to represent such a case. Yet such strange plants as Monophyllaea (fig. 6) and allied genera of the Gesneraceae living on the enlarged cotyle- dons might be examples. Cited literature: (1) Rec. Trav. Bot. Néerl. 1 (1904) 128. (2) De Trop. Natuur 1936, Jub. uitg. p. 73. (3) Now described as a separate species Canangium fruticosum CrRatB (Kew Bull. 1922, p. 166) being cultivated in Siam. (4) Compare F. A. McCLure, in Lingn. Sci. Journ. 12 (1933) Suppl. p. 119-149. (5) Philip. Journ. Sci. 74 (1941) 153-156. (6) Gard. Bull. Str. Settlem. 5 (1932) 245 seq.; ibid. 9 (1937) 139 seg. (7) Jugendformen und Bliitenreife im Pflanzenreich 1905. (8) cf. COCKAYNE, 13th Meeting Australas. Ass. Adv. Sci. (1912) 217 seq. 3. Dimorphous foliage It was observed by F. W. WENT (1) that in trees generally the foliage of the lower branches is larger than that of the upper twigs. He ascribes this to the amount of water available to different shoots (internal water-conducting capacity) ; so, in mature trees the upper foliage would be insufficiently pro- vided with water. The leaves of water sprouts, on the other hand, are mostly exceedingly large as their water supply is abundant. Leaves of these shoots are mostly hardly recognizable in the her- barium, as they may reach disproportionate di- mensions. Foliage for description in the herbarium ought therefore to be comparable and preferably that of flowering twigs. The dimorphy of the foliage is mostly linked up with a difference between flowering and non- flowering parts of the plant, similar to that found in Hedera. It is conspicuous in several climbing Ficus, Piper, Araceae, and in some Conifers. A strik- ing example of plagiotropically flowering twigs is that of Abroma angusta L.f. A good illustration is also Luvunga sarmentosa (BL.) Kurz (Rutaceae). The stem shoots of this Dec. 1948] General considerations XXII liana possess large straight axillary thorns and 1-foliolate leaves. The climbing shoots, however, possess conspicuously curved thorns and 3-foliolate leaves and the flowering parts of these are often unarmed. L. eleutherandra DALZ. was based on a type different from BLUuMe’s but is actually the same species, as was found by Kurz (2). Fig. 7. Heterophylly in Ficus deltoidea JACK (= F. diversifolia BL.) (Morac.), Mt Gedeh, W. Java, X 2/3. Putting into practice what he had read of Hedera helix in a botanical manual, Mr BoLtt made a remarkable application of the dimorphy of Piper cubeba L. Near Semarang, instead of cultivating it as a climber he took cuttings of the flowering twigs, and got shrublets which, though small, pro- duced abundantly ‘tail pepper’. Plants with dimorphous foliage are very numer- ous in Malaysia and species are frequently named after this peculiarity. Ficus deltoidea JACK (= Ficus diversifolia BL., fig. 7) is one of them; L. VAN DER Pu (3) could not find any regularity in its hetero- phylly. In Faradaya dimorpha PuLte from New Guinea there are two kinds of twigs, with decussate and with 3-whorled leaves of different shape. Phy- tocrene macrophylla BL. has both entire and 3-lobed leaves on one individual, as have Broussonetia sumatrana Miq., Knema heterophylla WARB., Ssever- al species of Gmelina and Sterculiaceae, Tiliaceae, Artocarpus varians Miq. A good case is also Uraria picta Desv. (fig. 8). Heterophylly is common in ferns. Polymorphy in leaf shape among different indi- viduals of a population is a subject which ought to have a separate heading. It is of universal oc- currence in the Malaysian flora, and has (e.g. in Cucurbitaceae) given rise to a multiplication of names. In Coccinea, MIQUEL (4) distinguished two species, one with incised leaves and one with angu- Jar entire leaves: according to BACKER they are identical, the incised leaves mostly belong to juve- Fig. 8. Uvaria picta Desy. (Legum.), with heterophyllous foliage, Kangean Island, moist Imperata fields at low alt., x 1/2. XXIV nile specimens. In Gymnopetalum cochinchinense (Lour.) Kurz there is even more confusion: speci- mens with incised leaves have been described as G. septemlobum Mia., G. quinquelobum Mig. and G. quinquelobatum CoGNn., those with angular or entire leaves as G. piperifolia Miq. and G. hors- fieldii Mia. There is probably a host of other names Soe HO XO) Bi, 2 PEER Nie o» , } 4 > Nee ~~ Fig. 9. Macrobiocarpy in Callistemon speciosus DC. (Myrt.) with 3 sets of fruits below the terminal bud, originating from 3 previous flowering periods, « !/4. for this species which is very variable in foliage. In Trichosanthes a similar polymorphy of the foliage caused superfluous description of species. Tr. gran- diflora BL. is, according to BACKER, a form of Tr. globosa BL. with lobed leaves. In Gynostemma CLARKE and BACKER assume a variability in foliage (3-foliolate to pedately lobed leaves) which will cause a considerable reduction of the number of species. Similarly scores of superfluous names are found in polymorphic species such as Urena lobata FLORA MALESIANA [ser. I, vol. 4! L. etc. In the Oleaceae, Nyctanthes dentata BL. is only a dentate-leaved form of N. arbor-tristis L. In some Pteridophytes heterophyllous leaves are well known. The most striking examples occur in the genera Teratophyllum, Stenochlaena and Loma- riopsis where according to Hottrum (5) hetero- phylly has caused much taxonomic confusion. An other striking case is that of the plant which is mostly cited as Lindsaya repens (BORY) BEDD. as demonstrated by W. TROLL (6). Cited literature: (1) Handel. 5e Ned. Ind. Nat. Wet. Congres (1928) 385-392 (1929). (2) Journ. As. Soc. Beng. 39 (1870) 69. (3) De Trop. Natuur 27 (1938) 89. (4) Flora Ind. Batavae 1, 1 (1855) 673. (5) Gard. Bull. Str. Settlem. 5 (1932) 245; ibid. 9 (1937) 139. (6) Flora 126 (1932) 408. 4. Dimorphous seeds and fruits Of heterocarpy (1) only few examples are known in the Malaysian flora. In some Compositae the marginal fruits are sometimes strikingly different from those produced by the central tubular flowers, as was described for Synedrella nodiflora GAERTN. by A. Ernst (2). It is also known that in Tragia volubilis L. normal and 2-hooked one-seeded fruits may occur together. In Umbelliferae normal fruits and fruits with one half reduced may sometimes be observed. In Leguminosae also different types of fruit are sometimes found on one plant. In Desmodium heterocarpum DC. the lower pods are 1-seeded, the upper 5—7-seeded. Dimorphous fruits and seeds are known in Aeschynomene spp. and in the genus Jussieua. One of the most curious cases of dimorphous fruits is that detected by BACKER (3) in the common Acalypha indica L. in Java where the tip of the male spikelets is crowned by a single female flower de- veloping into a T-shaped fruit with a central fertile and 2 lateral sterile cells; the central cell seems to be sunken in the tip of the axis of the rachis. The normal capsule consists of 3 equal cocci. A special case is that of macrobiocarpy (4) when not all fruits dehisce at the end of the season but a number remain closed on the plant and grow for years larger and woody. Sometimes fruits of 3-4 seasons are found on one twig, which thus keeps a reserve of seeds. Macrobiocarpy seems to be mainly restricted to the semi-arid climates and is of definite advantage in fire-swept areas. It is very common in some genera of capsular Myrtaceae (fig. 9), viz Leptospermum, Eucalyptus, Melaleuca, Agonis, Metrosideros, Syncarpia. It possibly also occurs in some Proteaceae, Coniferae, Casuarina, and some Rubiaceae. The woody structure, large size and modified shape of the fruits formed in previous seasons must be allowed for in identifying the species. Inadequate material may cause considerable confusion. Cited literature: (1) DELPINO, Mem. R. Ac. Sc. Inst. Bologna V, 4 (1894). (2) Ber. Deutsch. Bot. Ges. 24 (1906) 450-459. (3) Onkruidflora Jay. Sui- ker. (1930) 406-407. (4) WINKLER, Ann. Jard. Bot. Btzg 20 (1905) 37-41. eae oe ry a i ee th ae, . a] Pa L December 1948 We have the pleasure to send you herewith for your information the first part of ‘FLORA MALESIANA’ series I: Spermatophyta (flowering plants). Flora Malesiana will be a long-range work on the Botany of the Malaysian Archipelago, including the Malay Peninsula, Indonesia, the Republic of the Philippines, and the island of New Guinea. The work will be accomplished through the combined efforts of numerous col- laborating scientists on an international basis of co-operation. We hope to have made clear—in this first instalment—the prospects of its progress and the technique of publication which is slightly different from that in similar floristic works, but which was adopted because of our ardent wish for a - rapid conclusion of the work. We are convinced that your institute, or you personally, will be interested in series I of this flora, which is estimated to comprise some 2400 genera of Phane- rogams and about 25.000—30.000 species. If it falls outside your interest we beg to forward your copy to befriended botanists or return it to us. We ask your attention for the reduction in price in case of subscriptions by collaborating and co-operating institutions: the Staffs of the former take an active share in the composition of the work, whilst the latter promise to give assistance through the loan of specimens, information about collections, bibliographical assistance, &c. A similar advantage is offered to individual co-operating botanists. For the general (introductory) volumes 1-3, a separate subscription is opened for the convenience of non-botanical institutions and libraries. We are quite satisfied e.g. that volume | gives an approximate key to the history of botanical exploration of this large part of the Southwest Pacific, and is, therefore, indispensable as a source of data for historians, geologists, zoologists, and geographers. The issue of these single volumes will be limited, and will be in proportion to the subscriptions. The price is, provisionally, fixed at one guilder Netherlands currency pro 16 printed pages, incl. postage. It is estimated that, averagely, each year 240 pages will be issued. B. SCHOUTE DIRECTOR NOORDHOFF-KOLFF N.Y. : ORA MALESIANA Series I. SPERMATOPHYTA VOLUME 1, 2, 3 OF SERIES I OF FLORA MALESIANA with reduction for a collaborating institute with reduction for a co-operating institute with reduction for a substantially collaborating monograph es ees } TO TO P.O. BOX 151 ‘HAM 54 (MASS,), U.S.A. Y | t ‘ i ) ; t ‘ we at yi tel a CAST Oe pel ot my oe od me et ae ft Dec. 1948] General considerations XXV 5. Dimorphous flowers A most peculiar case of flower dimorphy occurs in some Orchidaceae viz in Renanthera lowii RCHB. f. (1) and Grammatophyllum speciosum BL. (2). The shape and colour of the lower flowers in a raceme are very much different from those of the upper ones. In these Orchids the occurrence of aberrant lower flowers is a normal phenomenon. It seems also to occur in some species of Arachnis and less obviously in some species of Bulbophyllum. In Oberonia imbricata LINDL. the upper flowers of the spike are abnormal and their gynaecium is reduced. Dimorphous flowers are also frequently found in dioecious and polygamous plants. Male and female flowers are sometimes very different in shape and size, e.g. in Mangifera, Brucea, He- vea, &c. In thyrsoid inflorescences the marginal flowers are often different from the central ones, or some- times the central ones are reduced or deformed. A conspicuous instance is Mussaenda where some flowers of the inflorescences have one calyx lobe large and leafy. Other cases are found in Hy- drangea, Sambucus javanica REINW., some Umbel- liferae, some Araliaceae, e.g. Boerlagiodendron, and some Mimosaceae. Cited literature: (1) WINKLER, Ann. Jard. Bot. Btzg 20 (1906) 1. (2) CosTERUs, Dodonaea 6 (1894) 24. 6. Cleistogamous flowers Cleistogamous flowers occur frequently in the Ma- laysian flora. A general survey has hitherto not been compiled. They were described in Clitoria by HArms (1) and RANT (2) where they are sometimes more fre- quent than normal flowers. The description of specimens with cleistogamous flowers has led here to phytographical confusion: the American genus Martia LEAND. SACR. was based on acleistogamous leguminous plant which is, actually, according to BENTHAM and HArms (/.c.) nothing but the cleis- togamous state of Clitoria. Cleistogamy also occurs in Malaysian species of Viola. It is stated by BEccARI (3) to occur in several Bornean Annonaceae. A very typical example is described in Commelina benghalensis L. by J. VAN WELSEM (4): cleistoga- mous flowers are present on subterranean shoots. - Another well known case in a common plant is Ruellia tuberosa L. mentioned by VAN WELSEM (5) and A. F. G. Kerr (6). Cleistogamous (better: c/leistopetalous) flowers are common in Orchidaceae as J. J. SmirH and R. SCHLECHTER both frequently mentioned. The for- mer gives a list of cases known to him in connection with his experience on autogamy (8); the latter studied the occurrence of cleistogamy especially in New Guinea (7) and found it in several genera, and both in the lowland and in the mountains. Some- times in several specimens all the flowers are cleis- togamous, e.g. in Eria rugosa LINDL. and Dendro- bium gemellum LINDL. SMITH even found species which are only known in the cleistogamous state (8, p. 138), or of which normal flowers have only occasionally been found. SMITH suggests that cleistogamy is more common in the rainy season, and he mentions that R. SCHLECHTER also got the impression that cleisto- gamy was common in very wet places in Sumatra and in the mossy forests of New Guinea more frequent in the rainy season than in the dry period. The same phenomenon has been observed by C. A. BACKER (9) for cleistogamy in Dicliptera canescens NEES (Acanth.) in Java; in moist countries or dur- ing wet periods in the dry season this plant pro- duces minute white cleistogamous flowers the corollas of which drop in the early morning. It is certainly noteworthy that a single trivial character like cleistogamy can so upset taxono- mical judgment that a new genus has been based on this abnormal state of a plant; this character changes the whole floral development, and sup- presses the manifestation of numerous genom tendencies in the mature plant. Physiologically this can only be explained by some break in the physiological chain reactions in an early stage of the development of the flower. The field observa- tions mentioned above may show how this problem may be studied experimentally. Cited literature: (1) Ber. Deutsch. Bot. Ges. 25 (1907) 165-176. (2) Ann. Jard. Bot. Btzg 44 (1935) 239-242; Bull. Jard. Bot. Btzg III, 4 (1922) 241. (3) Wanderings in the great forests of Borneo (1904) 402. (4) De Trop. Natuur 4 (1915) 142; see also BACKER, Handb. Flora Java pt 3 (1924) 25. (5) De Trop. Natuur 2 (1913) 53-58, 68. (6) Journ. Siam Soc. Nat. Hist. Suppl. 10 (1935) 66-67. (7) Die Orchid. Deutsch Neu Guinea, FEDDE, Re- pert. Beih. 1 (1914) p. Li. (8) Natuurk. Tijdschr. Ned. Ind. 88 (1928) 122-140, Orch. Rev. 37 (1929) 75, Nova Guinea 14 (1929) 359. (9) Onkruidflora Jav. Suiker (1931) 676, in nota. Teratologo-morphosis 7. Teratological forms Malaysia can boast of a series of good articles by J. J. SmitrH & J. C. Costerus (1) dealing with teratological phenomena in plants. Though several of these teratological forms are due to some hereditary factor, others are apparently caused by external factors. Some are possibly the result of a fungus’s attack though no fungus has been found. Pometia pinnata Forst. almost always has pe- culair large brown structures like witches’ brooms by which the tree can easily be recognized in the riverine forest: they suggest inflorescences. Invirescentia are quite a common phenomenon in several Compositae (fig. 10); the fact that they are often found together in colonies in several different species suggests that they may be due to some virus (?). Monstrous flowers occur rather frequently in Orchidaceae and have often confused systematists. XXVI FLORA MALESIANA [ser. I, vol. 4! J. J. Smit (2) has given an interesting account of them. The absence of a rostellum is closely con- nected with autogamy. As a result the flowers often hardly open, do not develop well, and their colour is paler than normal e.g. in Phajus tankervilliae BL. Fig. 10. Invirescence of Emilia sonchifolia DC. (Comp.) from Mt Abang, Bali, sandy riverbed, ca 1000 m, X 1/2. Sumatran specimens are apparently more normal than Javan. Of quite a number of these abnormal Orchidaceae no normal specimen is as yet known. Another abnormality is a variation in the num- ber of anthers, which, in Dilochia pentandra RCHB. f., is five; this ‘species’ is, however, a mere form of D. wallichii LiNDL. In other cases the third stigma- tic lobe is changed into a rostellum and the ros- tellum has become a stigmatic lobe. J. J. SmirH remarks that the phenomenon of peloria occurs in different degrees. Mostly the peculiarities of the labellum disappear, sometimes the tepals show some characteristics of the labellum. As peloria is for the most part inherited these forms are treated in more detail in the following chapter, paragraph 2. Teratological aberrations frequently cause such large changes in the structure of flowers that they strongly suggest some taxonomic novelty. An ad- ditional example is: an interesting 3-seeded coco- nut (3). DE Wir & PosTHUMUS collected at Buiten- zorg, Sept. 1944, a specimen of Cassia mimosoides L. of which each flower possessed 2 ovaries. This character is considered to be primitive or ancient in the Leguminosae; it has been reported to occur in several Caesalpiniaceae, e.g. in African Schwart- zia (6) and Indian Caesalpinia (7). In Archidendron, a genus of Mimosaceae centred in New Guinea, it is a generic character. Monstrous forms occur frequently in ferns where the plasticity seems greater than in Spermatophyta. Forked, lobed, and crisped leaves occur in many genera. Sometimes these monstrosities seem to be inherited and of racial rank (4). Even precocious spore formation may be partly inherited. Teratological aberrations merge gradually into individual variations. It is questionable whether an individual of Cassia mimosoides L. with two ovaries is to be classed as a teratological or individual variation. I will mention only a few examples of individual variation. MELCHIOR found (5) some flowers in Aphania masakapu Metcu. with free anthers. BACKER found individuals of A/ysicarpus rugosus DC. with 2-3-foliolate leaves. The leaves of Cis- sampelos pareira L. are sometimes both peltate and non-peltate in one plant. Some specimens of Ama- ranthus spinosus L. are unarmed. There is no end to this kind of individual vari- ation which sometimes affects typically structural characters. Experiments are needed to ascertain whether these aberrant plants are sports of the genom and hereditary or not. Cited literature: (1) Ann. Jard. Bot. Btzg vols 13, 19, 23, 24, 28, 29, 32, 33, 39, 42 (1895-1931). (2) Natuurk. Tijdschr. Ned. Ind. 88 (1928) 122-140. (3) Natuurwet. Tijdschr. Ned. Ind. 101 (1941) 144. (4) O. PostHumus, De Trop. Natuur 25 (1936) 177-178. (5) Notizbl. Berl.-Dahl. 10 (1928) 277. (6) JACQUES-FELIX, Bull. Soc. Bot. Fr. 92 (1945) 158. (7) Wicut & Arnott, Prod. Fl. Pen. Ind. Or. (1834) 281. Hypselo-morphosis 8. Phenotypic effect of altitude G. Bonnier, and later F. E. CLEMENTS, experi- mented on the effect of altitude on plants. BONNIER even assumed that species might change under pro- longed exposure to different conditions into other species but it seems that his experiments are un- trustworthy (1). In the Malaysian mountains where collectors are often compelled to follow ridges, plants from exposed situations are frequently brought home. Their foliage is often reduced, the leaves roundish, margins recurved, texture coriaceous, venation prominent, petioles reduced, habit compact. It is Dec. 1948] General considerations XXVII not always certain that these characters are a ‘nor- mal feature’ of the species. It is, therefore, of the greatest importance to try to collect such species from less exposed habitats (light, wind, poor soil), i.e. from the more fertile, sheltered, though less Fig. 12. Histiopteris alte-alpinav. A.v.R.(Polypod.), an altitudinal form of H. incisa J.SM., in its habitat between ‘sterile’ rocks on the summit of Mt Kerintji, W.Sumatra, ca 3750 m alt. (FREY WIJSSLING) accessible slopes. Extensive notes and large col- lections may show that such variability exists and serve to define the position of transitional speci- mens. The same species may be a crooked gnarled shrub when growing on a ridge and a moderately tall tree 50 m lower on the slope. The dwarfing of trees towards the summits of mountains and ridges is chiefly due to the gradual disappearance of the bole with increasing altitude. This is partly a consequence of the development of the young plants under a gradually increasing light intensity which stimulates branching close to the base. I observed a striking example in the field of dwarfing in Casuarina junghuhniana MiQ. on Mt Soeket, Idjen volcano, E. Java. Herbs too are generally dwarfed at high altitude, e.g. Erigeron linifolius WILLD. I studied an instructive case of variation induced by altitude combined with poor rocky soil in the grass Isachne pangerangensis Z.M. (fig. 11). A large series of transitions from tall to dwarfed speci- mens were represented. An example of a ‘hypselo-morphosis’ which has been described as a local-endemic species is that of the fern Histiopteris alte-alpina v. A. vy. R. (fig. 12) from the summit of Mt Kerintji, West-Central Sumatra, which is found at ca 3700 m alt., on a barren rocky ridge. This is certainly only a form of the common volcanophile H. incisa J. SM. It is difficult, however, to single out the various factors associated with increasing altitude viz more wind, sudden and large changes of temperature, strong insolation, poorer soils, lower atmospheric Fig. 11. Variable habit of Isachne pangerangensis Z.M. (Gram.) in N. Sumatra, Mt Losir, a. on burnt ridge in thick humus, 1500 m, c. on ridge with ericoid scrub, half-shade, thinner soil, 2000 m, d-e. open sandy flats on poor soil, 3000 m, f. on rocky windswept summit, soil nearly absent, 3440 m, x !/4. XXVIII FLORA MALESIANA [ser. I, vol. 4! pressure, different fluctuations of atmospheric hu- midity, greater difference between day and night temperatures, &c. In the absence of experiments one can only make some suggestion, in many cases based on observation in the field only. I know of only few species which are hairier in the mountains than in the lowland, e.g. Hydrocotyle sibthorpioides LAMK, of which BLUME described the hairy form as H. hirsuta BL. nonal. However, glabrous forms of this species also occur on the mountains! An other example is that of Dodonaea viscosa (L.) JACQ. Kurz (2) in his ‘Sketch of the Vegetation of the Nicobar Islands’ has remarked on the apparent absence of any general relation between hairiness and environment. There is no general rule that flowers are brighter coloured in the mountains. Ageratum houstonia- num MUL. has larger capitules and brighter blue flowers in the mountains than at low altitude buton Mt Pakiwang, S. Sumatra, I found the reverse (3), Scutellaria javanica JUNGH. var. sumatrana BACKER having here blue flowers at the base of the peak but white ones towards the summit. Of Dendrobium jacobsonii J.J. S. (§ Pedilonum) from the Casuarina forests 2400-2900 m alt. in East Java, J. J. SmirH says that at Bandoeng at 700 m alt. cultivated specimens had smaller and paler coloured flowers with a slightly different flower shape: mentum not bent and differences in the labellum; the inflorescences were, moreover, sometimes 2-flowered (4). Fruiting and flowering are also strongly influ- enced by altitude, as I demonstrated elsewhere (5). Experiments on the influence of altitude, the morphological and physiological behaviour of Malaysian plants have been scarcely made. TEys- MANN made some observations in his pioneer work on Mt Gedeh in West Java but did not comment; CosTER (6) wrote a note on the beech specimen planted by the former. In the Malay Peninsula RIDLEY (7) made some notes on the acclimatization of plants and the ways in which they can be accomodated at low altitude. Cited literature: (1) The New Systematics 1940, p. 55 seq. (2) Journ. Asiat. Soc. Beng. new ser. pt II, 45 (1876) 126. (3) Bull. Jard. Bot. Btzg III, 13 (1933) 16. (4) Bull. Jard. Bot. Btzg II, no 26 (1918) 41. (5) Bull. Jard. Bot. Btzg III, 13 (1935) 331-343. (6) Ann. Jard. Bot. Btzg 35 (1926) 105. (7) Agric. Bull. Str. & Fed. Mal. St. volumes 6-7 (1907-08). Photo-morphosis 9. Epiphytes It is sometimes wrongly assumed that epiphytism is confined to specific plants which are restricted to this mode of life. The amount of light appears to be the main factor. On the floor of closed forest the shade prevents epiphytes from making use of patches of bare soil, which in primary forest are always present. Exposed places, such as rocks, lava streams, landslides, poor silicious soils, mud streams and solfatara, however, offer conditions suitable for their growth, and are indeed often the places where many epiphytes are assembled, i.e. selected from the neighbouring forest. Though epiphytes may withstand dry conditions well, they mostly need a rather high atmospheric humidity which, in these exposed places, becomes a limiting factor. Most astonishingly rich communities of epiphytes I found on the often misty slopes of Mt Télong in N. Sumatra which from 1800 m upwards is like a rock garden carpeted with normally epi- phytic orchids amidst luxuriant dripping cushions of hepatics and mosses with some isolated dwarf Rhododendrons. It is sometimes contended that these terrestrial epiphytes are epilithes but I have Fig. 13. Vaccinium laurifolium Mia. (Eric.) as a hemi-epiphyte, height ca 5 m, along a road above Trétés, 1500 m, N.slope of Mt Ardjoeno, E. Java. Dec. 1948] General considerations XXIX also found them in deep humic soil between the rocks. I did not succeed in detecting any essential differences in habit between terrestrial and epi- phytic specimens. SONY nia § ahi ANNI yan a ay Po TEQk AY A OW, aA Hs VAVAVA IIS aes SARA o eae EASA AA I ee >) AV EY ? eR Ly ANY iy Gee el & f ar pe . 4 My) Y y, Vi Fig. 14. Habitat variations of Gentiana quadrifaria BL. (Gent.) in Java. The condensed pin-cushion shape is found on open dry windswept habitats, the loose habit on marshy or slightly shady soil, x 1/2. mens may differ considerably in habit from epi- phytic; they become more rigid and condensed, often fastigiate. Vaccinium lucidum (BL.) Miq., as an epiphyte is a loosely and irregularly branched shrublet with a tuberous woody base. Terrestrial specimens on ridges are mostly cupressus-shaped miniature trees without the woody tuberous base. Similar differences are found in Ficus deltoidea JACK of which the epiphytic and terrestrial speci- mens may differ considerably in habit. It goes without saying that a proposal by NAKAI (2) to distinguish the Ericaceous Agapetes and Vac- cinium by a terrestrial habit in the latter and an epiphytic habit in the former did not meet with the approval of SLEUMER. Some species begin their life as epiphytes but, when their roots subsequently reach the soil, they may grow into trees and sometimes show no sign of their early history. Such is found e.g. in Ficus, Fagraea, Schefflera, Wightia (1), and I even found it once in Vaccinium laurifolium Mita. (fig. 13). Many, however, are equally able to germinate ter- restrially and grow normally to trees. This is, in Wightia, even more common than the hemi-epi- phytic habit. As a small tree it is gregarious on the sunbaked lava streams of Mt Idjen in East Java, but on the forested outer slopes of the same moun- tain it is a hemi-epiphyte. Cited literature: (1) Revision of Wightia, Bull. Jard. Bot. Btzg III, 18 (1948) in the press. (2) Japan. Journ. Bot. 12 (1936) 37-38. 10. Shade forms Shade forms are found both in the lowland and the mountains. In general they possess larger, thin- ner leaves, longer internodes, &c. Shade and nor- mal leaves may occur in one individual. A very good example is Gentiana laxicaulis Z.M. de- scribed from Java, which appears to be a shade form of G. quadrifaria BL. Sometimes compact tussocks of the latter bear on one side shoots of ‘Jaxicaulis’ in one individual plant (fig. 14). Slender modifications of herbs can be observed in tall grass fields, comparable with those in temper- ate corn fields. These weeds growing in the damp dark micro-climate between the closely set culms of Saccharum spontaneum L., Andropogon amboi- nicus (L.) MERR., efc. strive for light. They show reduced leaves and inflorescences in relation to their lank habit. All herbs unable to emerge from the tops of the grasses show a similar habit, a kind of etiolated growth combined with some degree of nanism. For the effect of light on the habit of forest trees see the paragraph on savannah trees. Hygro-morphosis 11. Influence of drought Hardly anything is known of the influence of drought, and the changes induced by it in the mor- phology and physiology of Malaysian plants. In Gerbera jamesonii BoLus I observed in the dry year 1945 at Buitenzorg an astonishing reduction in length of the peduncles in relation to leaf length. The size of the leaves was very much reduced dur- ing the same period in Turnera subulata So. (T. trioniflora AIv.). Similar behaviour is mentioned by BACKER (1) in Jatropha gossypifolia L. var. elegans M.A., a plant which is thoroughly naturalized in the dry regions of Java and the Lesser Sunda Islands; dur- ing the driest period of the dry season only minute, short-petiolate dark-brown leaves are produced. Flowering of some trees, e.g. Dipterocarpaceae, and probably bamboos coincides with unusually dry years. Higher fungi fructify after a dry spell. Cited literature: (1) Onkruidflora Jay. Suiker. (1930) 411. Hora-morphosis 12. Seasonal variation Seasonal variation as described in Europe (1) I have not found recorded from Malaysia. In the cultivated Hibiscus sabdariffa L. I have seen fruiting specimens flowering a second time; these flowers, however, were only half the normal size and, also, paler in colour. Field botanists should search for ‘autumn forms’ in periodically dry regions. XXX FLORA MALESIANA [ser. I, vol. 4! Seeds of seasonal plants germinating in the wrong season may sometimes grow into dwarfs. I ob- served such forms also in Hibiscus sabdariffa L. at Buitenzorg. These dwarfs were 10-15 cm high and Fig. 15. Oblique, wind-trimmed Tamarindus indica L. (Leg.) on a ridge at ca 600 m on Noesa Penida, SE of Bali Island (DE VooGpD) had 2-3 flowers producing good seeds; the flowers were mostly much smaller than those of specimens flowering in the optimal season. Of leaf-shedding trees flowers are often collected with immature foliage which may deviate consider- ably from mature leaves. A peculiar case is that Dichrocephala chrysanthemifolia’ a Fig. 16. Compact dwarfing of plants near the fumaroles on the summit of Mt Kembar, Ardjoeno, E. Java, 3100 m alt. Normal specimens left, dwarfed ones right, x 1/3. of some leguminous trees which produce leaves in flushes; the latter consist of pale or white or even pink-coloured limply hanging leaves which only slowly get their normal texture (Maniltoa). The distinction between annuals, biennials and perennials causes many difficulties in species growing both inside and outside the tropics, spe- cially when the duration of life is used as a charac- ter to establish taxonomic limits. I assume e.g. Centrolepis to be annual in N. Sumatra, though its perenniality in S. temperate regions is used as a distinctive generic character against allied genera. In some Gramineae species may be similarly vari- able, specially in tropical localities, and thus deviate from temperate representatives of the same species in a character which is, in grasses, generally assumed to be of importance for the delimitation of species if it runs parallel with other morpholo- gical differential characters. It is puzzling me how it is possible to interprete from herbarium speci- mens the duration of life of perennials flowering during their first year and collected in that state. The use of the duration of life as a character in keying out species must be limited to very clear cases based on wide experience. Cited literature: (1) R. v. WETTSTEIN, Unt. ti. d. Saison-Dimorphismus im Pflanzenreiche. Wien 1900. 42 pp. Anemo-morphosis 13. Windforms A peculiar aberrant habit in shrubs and trees can be caused by constant winds. I have described this from Noesa Penida and Bali (1) in Terminalia catappa L., Barringtonia asiatica (L.) KURZ, Calo- phyllum inophyllum L., Bischofia javanica BL., Ficus sp., and Tamarindus indica L. (fig. 15). Other more recent examples are Dodonaea viscosa JACQ. near the Wijnkoops Bay, S. Java, and plants from Pa- dang Bolak in N. Sumatra described by M. VAN DER VoorT (2). These plants possess sometimes a peculiar oblique condensed one-sided habit and always show a decreased leaf size apparently owing to desiccation of the buds. They are found both on seashores and inland. Cited literature: (1) De Trop. Natuur 26 (1937) 69-78, 14 fig. (2) De Trop. Natuur 28 (1939) 201-209. Edapho-morphosis 14. Fumarole plants I have described (1) very aberrant modifications from some mountain summits viz Mt Ardjoeno in East Java (2) and Mt Agoeng in Bali (3) at 2900-3000 m alt. Some common lowland weeds, have through chance dispersal by wandering pilgrims and/or by deer established themselves in the immediate neighbourhood of fumaroles. Owing to the heat and moisture emitted by the fumaroles they are able to grow at these high altitudes. They are very much reduced in size and in habit very condensed, and their leaves are very small (fig. 16). Without Dec. 1948] General considerations XXXI flowers their identification would be difficult. They live in what may be called ‘open air hothouses’ in the subalpine zone, and the altitude, insolation, &c. are doubtless the factors which have induced their Fig. 17. Dwarf of Pemphis acidula Forsrt. (Lythr.) in flower and fruit, seashore of Oedjoeng Koelon, W. Java, X 2/s. aberrant mode of growth and resulted in what seems to be an ‘alpine habit’. The species concerned were: Hyptis brevipes Porr., Dichrocephala chrysanthemifolia (BL.) DC., Lycopodium cernuum L., Emilia sonchifolia DC., Bidens pilosus L., Oldenlandia herbacea Roxs., _ Fimbristylis capillaris A. Gray, Lindernia crustacea F. v. M. Fig. 18. Full-grown specimen of Pemphis acidula Forst., NE. coast of P. Tioman, Mal. Peninsula. (CORNER) Cited literature: (1) The Gard. Bull. Str. Settlem. 9 (1935) 63-69. (2) De Trop. Natuur 23 (1934) 119-120. (3) De Trop. Natuur 25 (1936) 158-159. 15. Rock plants; calcareous and silicious soils Both rocks and silicious soils may bring about rather conspicuous changes of habit in some plants, apparently owing to the small amount of nutrients available. These modifications can occur either at low or high altitude. Ze Mh) ( Seirus aS 5b J Sy > Ce 7 0 Fig. 19. Flowering and fruiting dwarf of Lepto- spermum flavescens J.SM. (Myrt.) on dry sterile sands of Toba highlands, Central Sumatra, in a heath-like vegetation, * 1/2. Mr C. N. A. DE Voocp collected dwarf speci- mens of Pemphis acidula Forst. (fig. 17) on the rocky coast of SW. Java resembling subalpine ‘Spa- lierstraucher’; normally this littoral species is a bush or small tree (fig. 18). On the so-called ‘padangs’, the gravelly or sandy flats of various geological history which sometimes occupy large areas in Sumatra and Borneo, many species are dwarfed: Leptospermum flavescens SM. when growing under optimal conditions is a me- dium sized cedar-like tree (fig. 20); here it is a dwarf, 10-20 cm high, which flowers and fruits abundantly (fig. 19). Many other species behave similarly. If herbarium specimens are not provided with good field notes, a botanist who has never visited the tropics is of course confronted with a puzzle. He may even find some other slight charac- XXXII FLORA MALESIANA [ser. I, vol. 4! ter not known to occur in the normal population and may think that they represent a different species: in this way another ‘paper species’ is created. Residents in the tropics ought to experi- ment with seeds gathered from dwarf individuals. Abandoned mining grounds in Banka, Billiton and Borneo are rich in dwarf forms of the most diverse species, which flower precociously as very small individuals (1). flowers, an unusual character in the species (2). Though no experiments have been made it is likely to be an edaphical form only. Scores of dwarfed species, mostly of shrubs or small trees but also of herbs (e.g. Dianella nemorosa LAMK. f. nana SCHLITTL. from Camarines and f. monophylla ScHuiTTL. from New Guinea) occur in the Philippine Islands, and especially in New Guinea on ridges in the mossy forest and the sub- Fig. 20. Full-grown specimens of Leptospermum flavescens J.SM. (Myrt.) on the slopes of Mt Bonthain, SW. Celebes. (L. VAN DER PIJL) On poor unweathered volcanic ash on the slopes of mountains the vegetation as a whole is dwarfed, e.g. on the slopes of the easily accessible Mt La- mongan, E.Java. Here the black gravel and sand is continually rejuvenated and gradually runs down. On these ash slopes all the common Javan mid- mountain trees and shrubs are dwarfed but flower and fruit profusely e.g. Radermachera gigantea (BL.) MiaQ., Parasponia parviflora MiQ., and Wein- mannia blumei (BL.) PLANCH., &c. flower and fruit on 1-2 m high shrubs. This observation induces me to suspect that Radermachera brachybotrys Me_rr. from Leyte merely represents a dwarf speci- men of some other species; KorTHALS found a similar specimen in the padangs of Borneo. I have also found dwarfs on wooded limestone cliffs in NW. Bali at 100-200 m mostly of her- baceous species. One of them was so aberrant that I described it as a new variety, Anisomeles indica (L.) O.K. var. biflora STEEN.; this had solitary alpine zone. No experiments have been done and the ‘normal’ habit of these plants is thus unknown. On rock cones, e.g. Mt Idjen, E. Java (fig. 21) and Mt Agoeng, Bali, I found Casuarina (fig. 22), Vaccinium, Rhododendronas extremely small shrubs and ascribed this to the very poor soil, though on these cones the influence of climate and soil are not readily separable. W. TROLL found precocious spore formation in Gleichenia vulcanica BL. on Mt Gedeh. This was certainly not caused by altitude but by the locally poor rocky soil. In cracks of rock on the summit Argapoera, of Mt Jang, E.Java, I have collected microphyllous specimens of a Polygonum which I originally took for P. chinense L. but which DANSER afterwards identified as an aberrant form of P. runcinatum Down (fig. 23). On Mt Kerintji were found minute fruiting specimens of Aralia ferox BL. which I have distinguished as f. nana (3). Dec. 1948] General considerations XXXII Cited literature: (1) TEYSMANN, Nat. Tijdschr. Ned. Indié 32 (1873) 84; DUNSELMAN, De Trop. Natuur 27 (1938) 97-104. (2) Bull. Jard. Bot. Btzg III, 17 (1948) 389. (3) Bull. Jard. Bot. Btzg LI, 17 (1948) 394. 16. Solfatara plants Specimens collected in craters are often of asurpris- ingly dwarfed habit even when old. SCHRGTER (1) figured a dwarf plant of Vaccinium varingifolium Mia. of East Java which was probably 50 years old and had the appearance of some alpine ‘Spalier- strauch’. At a short distance from these strongly insolated, edaphically dry and often wind-swept barren rocky places on slopes or summits, the same species occurs in hollows or other sheltered places as well-developed shrubs or small trees. The dwarf shrubs of craters are often wholly appressed to the soil (with rooting branches!), with a matted and prostrate habit. Owing to the poisonous gases emitted by the solfatara or effect of the wind on ridges, their surfaces are flat and look as if clipped Fig. 21. Vaccinium varingifolium Mia. (Eric.) as poor prostrate shrubs (‘Spalierstraucher’) near Kawah Idjen, E. Java, ca 2000 m alt. Exceedingly poor, eroded, very young volcanic soil. This species also grows in the mountain forest on the ridge behind in ca 3-6 m tall trees. or pruned (2) (fig. 21, 24). The solfatara may shift its outlet and so release these plants from its influence: I found some partly grown into a fresh bush, proving that the plant had recently escaped from the reach of the gases, the prostrate section being the oldest part. The reverse may also occur; erect shrubs may be affected later by crater gases (3) which makes them one-sided (fig. 25). Fig. 22. Casuarina junghuhniana Mia. (Casuar.). Old dwarf from the summit of Mt Agoeng, Bali Island, 3100 m, ona rocky windswept cone, x 1/2. Solfatara plants thus represent forms different in habit, and herbarium botanists must handle these materials cautiously. In a general sense the ‘poor soil’-conditions cause nanism. In Malaysia adaptability to habitat and varia- bility in habit under extreme conditions is far greater than the average herbarium botanist sus- pects. It is difficult to interpret aberrant specimens from remote regions without a thorough field knowledge. Unfortunately this has led to the des- cription of many ‘paper species’ which may seem specifically distinct but, when studied under various natural environments appear gradually to merge in the range of modifications existing in many Linnean populations. XXXIV FLORA MALESIANA [ser. I, vol. 4! Cited literature: (1) Vierteljahrschr. Naturf. Ges. Ziirich 73 (1928) 584. (2) De Trop. Natuur 24 (1935) 142-144, fig. 2-5. (3) SO2z, H2S, Cl2, &e. Hydro-morphosis 17. Water- and swamp plants Phenotypic variations comparable to those known in Europe are also known in the Malaysian flora. Jussieua repens L., when growing on muddy soil Fig. 23. Polygonum runcinatum Don. (Polygon.). Below: apex of a normal plant. Above: a very uncommon form of Mt Argapoera (Jang massif, E. Java) from clefts in rocks (St. 10960), x 2/s. Bh pant pT /y//'% be rear we Fig. 24. Sketches of prostrate clipped habit of shrubs in the crater of Mt Papandajan, W. Java, ca 2000 m, through the combined action of wind and sulphurous vapours from solfatara. Above: Vaccinium varingifolium Miq., below: Rhododen- dron retusum (BL.) BENN. (Eric.) (drawn after photographs). through the lowering of the water level, changes into a conspicuously different land form with minute hairy leaves and very condensed habit; it takes some time to recognize this land form (1). Land forms are also known in Potamogeton, e.g. from Sumatra, and in Utricularia. Of Neptunia plena Btu. a land form is known. BACKER suggests that the endemic N. javanica Mig., a terrestrial endemic species in Java, is merely a land form of the common N. oleracea Lour. (2). Limnophila sessiliflora Bi. and L. indica (L.) Druce, in shallow water, have deeply divided leaves below the surface with gradual transitions to lobed and toothed upper leaves above the water level. In very deep water pinnatifid leaves predomi- nate, but in marshy grass fields only pinnatisect to dentate leaves are present. The amount of aerenchyma is closely related to the depth of the water. Many swamp plants fail to flower in deep water, but come rapidly into bloom when the water recedes (Lemna, Blyxa, Pistia, Azolla, Salvinia, Marsilea, &c.). Swamp forest trees are also affected by the amount of water in the soil. A conical base to the trunk, so well known in Taxodium, is found fre- quently in other swamp species e.g. in Gluta renghas L., Alstonia spathulata Bu., &c. but, in non-inun- dated soils, the swollen base of the trunk is not or scarcely developed. The same is true of aerial roots at the base of the trunk. In deep swamps they may resemble the stilt roots of mangrove. Such roots may not de- velop in the same species when it is growing on dry land e.g. Acmena (Eugenia) operculata (ROXB.) Merr. & PERRY. Root production in these cases is doubtless a direct response to the habitat. Cited literature: (1) De Trop. Natuur 2 (1913) 83, fig. 3. (2) Schoolflora voor Java (1911) 428. Fig. 25. Oblique growth of Vaccinium varingifolium Mia. caused by sulphurous gases of the crater of Mt Papandajan, W. Java, ca 2000 m alt., ca 1 m high. Dec. 1948] General considerations XXXV Phyto-morphosis 18. Fungus and bacterial diseases, and symbiosis Malformations caused by fungi have led to some errors in identifying Malaysian plants. Loranthus maculatus BL. is, according to DANSER, the com- mon Dendrophthoe pentandra Mia. with a fungus on the leaves causing black spots (1). In specimens of Cassytha filiformis L. from New Guinea Dr HatusimA found some tetramerous Fig. 26. Pseudo-flowering of bamboo; galls caused by Epichloe treubii (Fungi). Bot. Gardens, Buitenzorg, Java. flowers with a central column marked by little pits. The slender inflorescence was glabrous and the rest of the plant hairy. It was evidently a malfor- mation caused by a Peziza-like Ascomycete. Root deformities caused by Cyanophyceae are found in Cycas, Gunnera, &c. Structures like witches’ broom are often found in bamboos, and often regarded as immature flow- ering parts. These pseudo-flowers are galls caused by a fungus (fig. 26). A curious malformation in Pilea trinervia WIGHT consisting of conspicuous swellings of the inter- nodes was described by Mrs WEBER VAN Bosse (2) and is caused by a parasitic alga: Phytophysa treubii W. Vv. B. In Pavetta, bacteria cause dark often thickened spots in the leaves. According to BREMEKAMP the symbiosis is mostly restricted to particular species. Similar bacteria are found in species of Psychotria, Ardisia, the tips of the leaves of Smilax, &c. The presence of absence of bacterial nodules is used in the identification of Rubiaceae, a rather singular method. Cited literature: (1) Compare BOoEDIJN, Bull. Jard. Bot. Btzg III, 13 (1935) 497-501, fig. 1. (2) Ann. Jard. Bot. Btzg 8 (1890) 165-186. Zoo-morphosis 19. Ant plants (myrmeco-morphosis) Several Malaysian plants are inhabited by ants. TREUB (1) proved that the cavities in which the ants live in the tubers of Myrmecodia and Hydno- phytum are also formed in the absence of ants. In other instances, however, ants presumably bite their way into internodes and remove and carry away the pith. This was shown to occur in Endospermum moluccanum T. & B. (E. formicarum Becc.) by DocTERS VAN LEEUWEN (2); I am able to confirm this. DoCTERS VAN LEEUWEN also found some specimens uninhabited by ants. In Endosper- mum, therefore, whether the internodes are hollow or not is certainly not a good specific distinction, hough used by PAx in his key to the species of Fig. 27. Above: Kibessia sessilis BL. (Melast.) being based on a galled swollen fruit of K. azurea BL. (W. Java). Below: peculiar galls of Styrax benzoin DRYAND. (Styrac.) from Sumatra, X 2/3. XXXVI FLORA MALESIANA [ser. I, vol. 4! Endospermum subg. Capellenia (3). Moreover, as the other character used by PAx, viz the number of cocci in the fruit, varies from 3—5, through igno- rance of data on points the collector ought to have noted on the label, the whole key breaks down. In Wightia borneensis Hook. f. some individuals are attacked by ants which remove the pith from the upper internodes. The withdrawal of the inner tissue causes the hollow internodes to assume a cigar-like shape. Cited literature: (1) Ann. Jard. Bot. Btzg 3 (1883) 129-153. (2) Treubia 10 (1929) 1-7. (3) Pfl. Reich Heft 52 (1912) 34. 20. Galls deceptive to phytographers (cecidio-morphosis) Cecidia caused by animals have sometimes de- ceived botanists when describing plants. An ex- ample is Ceratostachys arborea BL., a genus based on a galled fruit of Nyssa javanica (BL.) WANG. Kibessia sessilis BL. is merely the galled and enlarged fruit of K. azurea BL. (fig. 27). According to RIDLEY (1) Apteron lanceolatum Kurz, described as a distinct genus, is identical with Ventilago kurzii R1IDL.; KURZ mistook some insect galls for the ripe fruit. MIQUEL described (2) an abnormal tree from Sumatra which was actually a species of Styrax, a genus in which most peculiar galls (fig. 27) are very common. Otopetalum micranthum Mia. is an Apocynacea described from Java. According to BOERLAGE (3) the plant was referred to the wrong tribe because MIQUEL erroneously took galled flowers for 1- seeded berries; the former author suspects that it is related to Micrechites. Insects (mostly cicads and larvae of Hemiptera) cause a singular malformation of the flowers in some species of the genus Sterculia. H. C. CAM- MERLOHER (5) observed that they are attracted to so-called ‘sugar hairs’ which occur on the inside of the perianth. The insects injure both the hairs and the outer tissue in an early stage of develop- ment of the flower. The calyx becomes enlarged, thicker and tough, and opens hardly in anthesis; its lobes remain short and triangular, and the tube is relatively large. These flowers are conspicuously different from the normal ‘uninhabited’ flowers and, according to ADELBERT (6), ought not to be used when describing or identifying plants. BACKER (7) described in Hibiscus schizopetalus (Mast.) Hook. f. malformations of the vegetative parts and of the flowers caused by plant lice. The Philippine species Euphoria malaanonan was described by BLANCO and by him referred to Sapin- daceae but MERRILL stated (8) that it is merely based on specimens of the echinate galls of Shorea guiso BL. of the Dipterocarpaceae. W. M. DocTers VAN LEEUWEN has published (4) an illustrated book on zoocecidia of Indonesia. Cited literature: (1) Flora of the Malay Peninsula 5 (1925) 300. (2) Linnaea 26 (1853) 285. (3) Hand- leiding Fl. Ned. Ind. 2? (1899) 380. (4) The Zooce- cidia of the Netherlands Indies, Batavia 1926; Supplement, Ned. Kruidk. Arch. 51 (1941) 122- 251. (5) De Trop. Natuur 22 (1923) 147. (6) In BACKER, Flora van Java, Nooduitg. [VB (1944) fam. 107, p. 18. (7) Flora van Java, Nooduitg. IVc (1943) fam. 109, p. 27. (8) Spec. Blanc. (1918) 33. 21. Influence of browsing animals (pascuo-morphosis) In some parts of Malaysia browsing cattle and deer (1) can induce changes in the morphology of plants which might be termed pascuo-morphosis. In Fig. 28. Casuarina junghuhniana Mig. (Casuar.) on Mt Jang, E. Java; crown trimmed below by deer. the deerpark of the Buitenzorg Palace, deer regu- larly feed on the pendent air roots of Ficus and prevent them from reaching the soil. The trees therefore remain single-stemmed and do not form thickets of pillar-like roots. A similar effect is caused by deer on Mt Jang, in East Java, where deer eat the hanging branches of Casuarina junghuhniana Mia. as high as they can reach (fig. 28). The trees look as if clipped (2) at the underside of the crown like those on the lawns at Buitenzorg. Much the same was observed on Mt Rindjani, Lombok Island (3). The broom- like appearance of the grass Pogonatherum pani- ceum HAcK. on Mt Diéng was ascribed to grazing cattle by the late Mr LOOGEN (4), an excellent ama- teur field botanist. Browsing of animals has in general the same effect on the vegetation as frequent burning: the plants acquire a low habit and flower at an early age (pseudo-nanism). This occurs very commonly on the closely cropped fields of fine grass of Mt Dec. 1948] General considerations XXXVII Jang. It is only in hedges, on steep slopes or some- where out of reach of deer that plants grow to normal dimensions. In Central and East Java, and the Lesser Sunda Islands pascuo-morphosis is due to browsing of cattle in the dry season; it is known in Zizyphus jujuba L., Streblus asper L. and other shrubs. These assume a fastigiate habit as high as the browsing animals can reach: above about 2 m the twigs are again spreading and form a globular crown. A transition to anthropo-morphosis is the clip- ping and pruning of plants which can sometimes produce an aberrant habit. According to BACKER (5) repeated cutting of plants along roadsides pro- duced a peculiar table-shaped densely branched SES AAAS ) A> a \ ay A) P ys S 3 (3 aa ee \ ae KS Wy) SR for samp CX) Es Ly ae ? AY WIS Sd Fig. 29. Seedling of Butea monosperma TAuB. (Leg.) sprouting in its 3rd year in the fire-swept savannahs of Indramajoe, W.Java, and developing a lignotuber, x !/3. form in Sida retusa L. near Batavia. In the Lesser Sunda Islands cattle are sometimes fed in the dry season with leaves of trees, as is done in Africa. For this purpose the people lop the lower branches of trees in order that cattle may reach the foliage. This causes a tendency to umbrella-shaped trees. Deer may cause the same change of habit. Cited literature: (1) Mentioned for India by HooKEeR & THOMSON, Flora Indica (1855) 29. (2) De Trop. Natuur 21 (1932) 27. (3) De Trop. Natuur 30 (1941) 123. (4) De Trop. Natuur 30 (1941) 70. (5) Flora van Batavia (1907) 102. Anthropo-morphosis 22. Influence of fire (pyro-morphosis) The changes in habit and structural characters induced by fire were named pyro-morphosis by PERRIER DE LA BATHIE who made observations in the island of Madagascar. In Malaysia there are few reliable data. One of the changes induced by regular burning of the vegetation is that plants are stunted, and flower when small. Owing to the damage done to the surface part of the plant the underground parts thicken, and the upper portion of the rootsystem and lower por- tion of the stem form gradually a thickened half- subterranean ‘lignotuber’, which sprouts after fires have swept the plains. I made some observations in the Indramajoe plains (W. Java), where species of Grewia, Butea, Dillenia, Morinda, Phyllanthus emblica, Zizyphus, &c. sprouted from _ these thickened bases (1) (fig. 29). Some instances of phytographical importance have come to my knowledge. RANT found (2, 3) that Psidium cujavillus Bur. f. can originate spon- Fig. 30. Park-like savannah in SW. Soembawa; trees with short boles and rounded crowns. (DE VooGcD) XXXVIII FLORA MALESIANA [ser. I, vol. 4! taneously from root shoots of Ps. guajava L. It is distinguished from Ps. guajava L. mostly by differ- ences of size. Ps. cujavillus BURM.f. must, therefore, be reduced to a sport of Ps. guajava. Fire-resistant trees are often crippled beyond recognition. Plants described from semi-arid (4) regions where fires occur annually are sometimes known only in this crippled state, e.g. Fordia fruti- cosa CRrRats, from N. Siam, described (5) as a shrublet 40 cm tall. The late A. F. G. KERR, a most able and experienced field botanist, stressed in a note made in the field that the plant was growing in an area subject to fire; this may explain such a habit in this otherwise arboreal genus. The normal plant will, in all probability, prove to be a tree. Cited literature: (1) De Trop. Natuur 25 (1936) Jub. nummer, p. 117-118. (2) Ann. Jard. Bot. Btzg 41 (1930) 27-32. (3) Natuurk. Tijdschr. Ned. Ind. 94 (1934) 112. (4) That is: regions which are periodically wet and dry, and show two distinct seasons coinciding with the monsoons. (5) Kew Bull. 1927, 60; Fl. Siam. En. I (1928) 395. 23. Pioneer plants In the preceding paragraphs 9, 14-16, and 22, several examples have already been given of pioneer plants. I am decidedly of the opinion that this term should not be restricted to plants peculiar to land- slides and other bare soils. Trees settling in savan- nahs or devastated areas, epiphytes settling on rocks, &c. are just as well ‘pioneer plants’. They constitute seral vegetation types. In the initial stages of revegetation forest trees may appear as pioneer shrubs, flowering and fruiting early. It is rather baffling to find Schima noronhae REINW. flowering and fruiting as a lax shrub 2m tall, when one is familiar with the gigantic full grown tree in the forest (height sometimes over 50 m, columnar bole over 1 m diam.). BACKER found (1) near Batavia flowering specimens 15cm tall of Grewia microcos L., usually a tree up to 17 m high. The same phenomenon can occur in Adinandra, various Urticaceae, Leguminosae, Ulmaceae, &c. In New Guinea some endemic species were originally described as tall trees but Mr Brass has recently found them as gregarious pioneer shrubs in differ- ent seral vegetation types. A promising shade plant, AJ/bizzia sumatrana STEEN., described from the Westcoast of Sumatra, was found to be a forest tree. In the plantations it is a weed tree flowering and fruiting at a very early age. In the forest, its native habitat, flowering is apparently suppressed by the deep shade, the seedlings grow into pole trees, slender and tall with a minute crown and real growth only starts when the crownlet emerges from the canopy. The behaviour and appearance of forest trees when growing in the open cannot be predicted. The adaptive capabilities of most Malaysian forest trees are unknown, though data on these points would be of great importance for practical forestry. See also chapt. 2 on precocious flowering, a phenom- enon frequently observed in pioneer plants. Cited literature: (1) Flora van Batavia (1907) 196. 24. Savannah trees Trees of the savannah generally differ in habit from trees of the closed forest in their short bole and spherical crown (fig. 30). If young forest trees Fig. 31. Altingia excelsa Nor. (Hamam.) at Tjibodas, W. Java, ca 1450 m. Left: forest-grown tree, clear bole ca 25 m. Right: planted on the lawns of the mountain garden, at ca 200 m distance. Dec. 1948] General considerations XXXIX in deforested areas become exposed, or when they are planted as roadside trees, they acquire this shape: a striking example is Altingia excelsa Nor., a forest giant of the West Java midmountain forest between 600-1600 m. The clear bole is usually a characteristic feature of the tree; it is columnar, up to 14m in diameter, and up to 20-30 m from the ground unbranched (fig. 31). Specimens 50 years old, however, planted on the lawns at Tji- bodas mountain garden, have grown into low spreading trees with hardly any bole at all (fig. 31). Descriptions of the habit of trees and shrubs taken from specimens grown in private and botanic gar- dens, will therefore generally not agree with those taken from specimens in the forest. THORENAAR made similar observations in Javan oaks, and other trees such as Podocarpus imbricata BL., Quercus, etc. The habit of trees grown in open gardens resembles the shape of trees of the savannahs where the rounded crowns on a short bole often charac- terize the physiognomy of the open savannah for- est. Physiologically this tree form is in all proba- bility determined by the high amount of light pres- ent during the juvenile stages of growth. The crowns of mature forest giants exposed after de- forestation also tend to become rounded. VARPATIONS*BOUND TO THE GENOTYPE (Genotypic variation) 1. General remarks 3 : XXX1X 2. Peloric flowers . , : xli 3. Other teratological or deviating forms xlii 4. Distribution of the sexes. xlili 5. Geographical segregation of Malaysian specific populations . ‘ - xliv (a) Ecological . xlvi (b) Regional xlvil (c) Topographical ‘ xlix 6. The problem of speciation . ‘ ; li 7. The effect of isolation . ; lii 8. Centres of speciation in the Malaysian flora . : 5 : ; ; : : lii Variation bound to the genotype is intimately related to the species concept and to geographical distribution. The ideas advanced by J. D. HOOKER in his introductory essay to the ‘Flora Indica’ (1855) have gained in ‘philosophical’ importance through the progress of basic research in experi- mental taxonomy in the last decades in Europe and the United States, in particular by E. BAur, F. v. WETTSTEIN, N. H. NILsson, G. TURESSON, A. MUNTzING, O. WINGE, G. D. KARPECHENKO, B. H. DANsER, W. B. TURRILL, J. CLAUSEN, J. P. Lotsy, H. DE Vries, N. I. VAviLov, and many others. This work is summarized in several useful symposia and textbooks such as ‘The New System- atics’ edited by J. HuxLEy, CAIN’s ‘Foundations of Plant Geography’ and CLAUSEN’s c.s. ‘Experi- mental Studies on the Nature of Species’ (1945). It lies outside the scope of this essay to consider the various view points on these subjects. Some of them I have already touched on in a study of Malaysian mountain plants.! Now I will try to explain briefly some current views, including my own, and will illustrate them by examples taken from Malaysian botany. 1. General remarks Scientific names of species rest on the ‘International Rules of Nomenclature’ and depend on the identi- ty of the ‘type specimen’. This need not imply that only studies in which all type specimens have been (1) Bull. Jard. Bot. Btzg III, 13 (1935) 358-391. 9. Centres of generic development in Malaysia . : - lv 10. Local-endemic species and genera : Ivi 11. Parallel or homologous variation ? Ivii 12. Reticulate affinities. ‘ Ivili 13. Vicariism in the Malaysian flora . : lix 14. Adaptation and migration . : : Ix 15. Hybrids in the Malaysian flora . : 1xii 16. Polyembryony, parthenogenesis and apogamy . : : : : lxiv iene origin of native aliens F - 3 lxv 18. The origin of Malaysian cultigens Ixvi 19. Extinct plant groups in Malaysia. Ixvili examined, full synonymy is given, in which nomen- clature is in accordance with the Rules, and in which new species or other taxa have been de- scribed by careful Latin descriptions, are sound and durable. I know some excellent works in which nomenclature is neglected and synonymy is obso- lete but in which botanical distinction and de- scription are superb, and specific delimination is carefully drawn. Such books give the impression that the author is master of his art. I become more and more convinced that in the past two decades the care for outward appearance has come to take a too predominant share of the attention of some botanists who wrongly assumed that the examination of type specimens is the last word in real taxonomical research. It is sometimes not realized that type specimens often are only deficient, poor and miserably dried single plants chosen at random from billions of specimens growing in Nature in the past, the present and the future, which, together, according to the Linnean principle, compose the specific population. Very often these type specimens by no means represent the ‘average’ or ‘most common type’ of the population. The ‘z-typica’-distinction, therefore, has only nomen- clatural, i.e. administrative but no botanical value. It needs no comment that an up to date nomen- clature is a conditio sine qua non for any taxonomic work, and it will be tried to reach a high standard in this Flora. At the same time, however, is is hoped that the contributors will not be satisfied when writing formally correct revisions, but also carefully consider the status of both genera and XL FLORA MALESIANA [ser. I, vol. 4! species and the structural differences distinguishing them, not merely limiting themselves to the dis- tinction of taxa for reasons of convenience. Inadequacy of material and lack of field knowl- edge are both sources for the provisional dis- tinction of ‘species of convenience’, which mark, in tropical floras, as a rule the initial stage of ex- ploration. Whilst the inadequacy of material is an immo- vable obstacle, and not every revisor will have the privilege to acquire field knowledge, a large fund of experience has been collected on the subject of variation in tropical plants. The following pages contain a discussion, valu- ation, and illustration of variation in Malaysian plants. In general the new systematics, based on modern experimental taxonomy holds that a narrow spe- cies concept is not in accordance with the structure of nature. Modern insight offers no support to so-called ‘splitters’, one of whom declared to me as his ‘prin- ciple’ that he felt obliged to distinguish the smallest distinguishable entities and to assign to these a binomium. On the average the standard of the specific concept proposed by LINNAEUS in his works and rules cannot be questioned and for binomiums the Linnean canon has priority of conception. I fail to understand how that conscientious splitter who in matters of nomenclature adheres strictly to priority and applies the binomial system of LIN- NAEUS can simultaneously call wide-spread poly- morphic populations ‘collective species’. To do so is intentionally to depreciate the time-honoured and scientifically sound Linnean standards. The difficulties confronting the systematist are manifold; no clue exists to the causes of polymor- phy. Systematists are still far from being able to explain why some species are polymorphic and variable and why others show a narrow amplitude of divergency. This is a fundamental barrier to the methods of ‘weighing and measuring’, a common basis in the natural sciences generally. Polymorphy is apparently not related to speciation, as mono- specific units such as Homo sapiens, Cocos nucifera, &c. are very variable and species of large genera are sometimes not very polymorphic, though it will be observed that in most genera at least one species is widely distributed and rather polymor- phic. Ignorant of the laws underlying his taxonomic distinctions, the systematist should be aware that he deals with unequal entities, though we may try to shape them as consistently as possible. A basic research in this connection is the work of the late E. BAurR, the geneticist, who made un- surpassed long-range efforts in the combined fields of taxonomy, field-work and experiments, to disen- tangle the genus Antirrhinum sect. Antirrhinastrum (1). This section had some dozens of local species described from the West Mediterranean distin- guished by characters which were the despair of taxonomists. BAUR proved that the population of the section falls apart in numerous ‘colonies’ or ‘partial populations’ which are isolated and cross mutually (convivia, sensu DANSER). Each colony has its own type, the larger the colony the wider the local diversity (fig. 33c). All these local types can be freely intercrossed with fertile offspring, and there is no doubt that, if not isolated in nature, they would together merge into a still more diverse popu- lation with transitions and intermediates. BAUR, moreover, obtained experimentally many forms not realized in nature (/.c. p. 289), showing that the potential variability (2) or polymorphy (3), i.e. the total number of possible forms (genetic capabilities) is not exhaustively represented. There is a great reserve of possible combinations, and the genus manifests itself to us at present under a limited number of combinations. Hardly any plant species is evenly spread within its area, and many occur in aggregates or colonies. BaAur’s findings are thus of the utmost value for the Malaysian archipelago where isolation is a normal factor in specific populations, in the low- land owing to the insular discontinuous nature of this region and in the mountains still more accen- tuated by the often long distances between the summits. We cannot expect that the whole plant world will be subjected to accurate and thorough experimental research like BAuR’s Antirrhinum- studies, but, judging from his results, it appears that geographical distribution is an important argu- ment when determining the status of taxa. Well- defined allied species possess in general overlap- ping areas of distribution proving the independence of the populations. If, however, several allied species exclude each other geographically one must be on the alert, and check the differential charac- ters again because the specific population may well be differentiated into a number of races, subspecies, or ecotypes. Especially along the frontier—horizontal and altitudinal—of the area, a species population has a different facies from that at its centre. KERNER (4) showed that in Cytisus sect. Tubocystis aberrant forms occurred along the border of the area (fig. 33e). Migrating plant individuals (seeds, spores, fruits, root-stocks, &c.) carry only a part of the potential polymorphy of the genus, and their off- spring will possess a special facies. Hence, along the frontiers, combinations can be expected which are not realized within the centre of the population. According to VAVILOV such pioneer aggregates are found to be recessive homozygotous; this is of great practical importance. It is worthy of note, as was pointed out by E. C. ANDREWS (5) that BENTHAM realized long ago that the geographical station of a waif or colonist im- poses variations upon it almost from the moment of its arrival. ANDREws adds, that Eucalypts plant- ed in New Zealand, California, etc. present marked differences in general appearance from the same species in Australia. Stimulated apparently by HooKeER (6), HUGO DE Vries (7), the Master of experimental taxono- mic botany, remarked that the initial stages of new species will be found most easily in luxuriant alien vegetations. His classic example, Oenothera, show- ed these ‘mutations’ — which they indeed are if the TAXONOMIC REVISIONS ACERACEAE'! (S. Bloembergen, Buitenzorg) 1. ACER mine, Sp.P). (1753) 1054; Pax, Pfl.R. 8 (1901) 17K. & V. Bydr. 9 (1903) 252. Trees or shrubs, buds with many perules. Leaves decussate, petiolate, entire, palmate or pinnate, appearing simultaneously with the flowers or later, exstipulate. Inflorescence racemose, corymbose or spicate, terminal with 2-4 leaves, or rarely terminal or axillary without leaves. Monoecious or dioecious, flowers actinomor- phic, d and 9, ovary in the d fls more reduced than stamens in 9 fis. Calyx and corolla 4-5-merous. Stamens 4-10, mostly 8, hypogynous or perigynous. Disc extra- or intrastaminal. Ovary superior, 2-celled, laterally flattened, each cell with 2 ovules. Fruit a samara, splitting into 2, rarely 3, winged usually 1-seeded parts. Seed without endosperm, radicle elongate, cotyledons foliaceous, or thickened, plicate, involute or flat. Distr. Ca 200 spp. in the N. hemisphere, only in Malaysia crossing the equator. Notes. By BLUME, BENTHAM & HOOKER, MIQUEL, &c. this genus was included in the Sapindaceae. In Malaysia only one species. 1. Acer niveum BL. Rumphia 3 (1847) 193; Pax, Bot. Jahrb. 6 (1885) 293; ibid. 7 (1886) 207, cum var. cassiaefolium; WWESMAEL, Bull. Soc. Bot. Belg. 29 (1890) 41, cum var. praec.; SCHWERIN, Gartenfl. 42 (1893) 228, cum var. laurinum & praec.; PAx, in E. & P. 3, 5 (1896) 267, 271; Pfl.R. 8 (1901) 4, 31; K. &. V. Bijdr. 9 (1903) 254; Backer, Schoolfl. (1911) 272; HeEYNE, Nutt. Pl. (1927) 987; Koorp. FI. Tjib. 2 (1923) 153; Merr. En. Philip. 2 (1923) 493; STEEN. Bull. J.B.B. m1, 13 (1936) 148.—-A. java- nicum (non Burm. f., 1768) JUNGH. Monatsber. Berl. Geogr. Ges. 1842; JUNGH. & DE VR. Tijd. Nat. Gesch. & Phys. 10 (1843) 138.—A. /aurinum HAssk. ib. nomen; Miq. Fl. Ind. Bat. 1, 2 (1859) 582, Suppl. (1860) 200, 511; BorrL. Handl. 1 (1890) 281.—Laurus alba BL. Rumphia 3 (1847) 193, in syn.—A. cassiaefolium BU. l.c.—A. philip- pinum MerrR. Gov. Lab. Publ. 35 (1906) 36.—A. curranii MERR. Philip. J. Sc. 4 (1909) Bot. 285.— Fig. 1. Tree up to 48 m, clear bole up to 28 m by 70 cm, buttresses to 2m high. At the start of the dry season foliate twigs sprout 2-4 together simultane- ously from last year’s buds; inflor. appearing in the axils of fallen leaves. Both flower and shoot- buds ca 4 mm long with 4-9 pairs of decussate ca 2mm long caducous perules. Leaves simple entire glabrous, glossy dark green above, glaucous, whitish or light blue-grey below; petiole 11/4-10 cm; blade elliptic to lanceolate, 3- to slightly 5-plinerv- ous at the base, apex acuminate to tailed, tip subacute. Inflor. corymbose, either d or 9, rarely with few fis of the other sex, glabrous, 2!/2-10 (in fruit to 19) cm long; peduncle !/2—-3!/2 cm, pedicels 4-17 mm. Flowers pale yellowish. Sepals and petals (3-)5, free, resp. 2!/2-3 and 1!/2~-2!/2 mm long. Stamens (4-)6(—8) in 1 whorl, sometimes isomerous and then alternating with the petals, attached on the disc in pits; filament in d 5 mm, in 9 2.2 mm; anther (1) In Malaysia only one genus. 3/4 mm (in Q slightly smaller and not dehiscent). Disc flat, glabrous to woolly. Ovary 2 mm broad, densely woolly, in od (with the styles) usually mor HS Fig. 1. Acer niveum BL. X 1/3, flower enlarged. strongly reduced. Styles 2, 1'/2 mm long. Wings of fruit 4-7 by 1-21/2 cm, asymmetric and obovate, inside narrowed or straight, hairy; mericarp proper 8-13 mm long, ovate. 4 FLORA MALESIANA [ser. I, vol. 41, Dec. 1948] Distr. Cf. fig. 2; in the Malay Peninsula re- cently collected in the hills near Cameron High- lands (CF. 27181, 27344, 36281, 37745, 45489), in W.Borneo once near Simpang at 27 m (bb 13518) and once in Sarawak (HAVILAND 2092), in the other islands many localities. Fig. 2. Distribution of Acer niveum BL.; in Borneo and the Malay Peninsula it is very rare. Ecol. In primary, rarely in secondary or devas- tated, forests, often common but scattered, 900- 2550 m, in Flores descending to 750 m, in Sumatra to 630 m, in Celebes to 450 m, and at only 27 m near Simpang, Borneo. Fls in April—Aug., fr. July—Nov. In Casuarina forest saplings have been found. When flowering the leafless crown swarms with Hymenoptera collecting honey. Trees are easily located in the forest by the fallen leaves which are glaucous and _ fine-reticulate-veined underneath. Vern. Some dozens of native names have been recorded, none of these fixed (HEYNE, /.c.). Uses. Timber unimportant, no distinct heart- wood is present; used for building purposes, fit for boxes. Notes. PAx inserted A. niveum in his sect. Integrifolia in which the inflor. is terminal on short leafy twigs. I could examine this character in A. oblongum WALL. (WALL. 1222 A), A. laevigatum WALL. (THOMSON s.n.) and ‘A. niveum’ (HELFER s.n.) from India. A. niveum BL. with its axillary leafless inflorescences is, however, possibly related to Pax’s sect. Lithocarpa and does not belong in sect. Integrifolia. Excluded Acer javanicum Burm. f. Fl. Ind. (1768) 221 = Actinophora fragrans R.BR. sec. BACKER in Herb. Bog.; according to BURRET it is Colona javanica, both Tiliaceae. PHILYDRACEAE (C. Skottsberg, Goteborg) Erect herbs with a short rhizome. Leaves linear radical or crowded at the stem base, distich, equitant, parallel-nerved. Flowers zygomorphic, bisexual, solitary in the axil of spathaceous bracts. Perianth corolline, segments 4, 2-seriate. Stamen 1, inserted at the base of the abaxial segments. Filament flattened; anther 2-celled; cells straight or twisted, opening lengthwise by slits. Ovary superior, 3-celled with axile placentas, or 1-celled with parietal placentas. Style simple. Ovules ~, ana- tropous. Capsule with 3 valves. Seeds ~. Distr. Centering in Australia, comprises 4 genera with 5 species. KEY TO THE GENERA 1. Philydrum 2. Helmholtzia 1. Outer tepals free. Anthers spirally twisted. Ovary 1-locular 1. Outer tepals united at the base. Anthers straight. Ovary 3-locular 1. PHILYDRUM BANKS & SOL. ex GAERTN. Fruct. 1 (1788) 62; Mig. Fl. Ind. Bat. 3 (1855) 250; Hassk. Bull. Soc. Bot. Fr. 16 (1869) xxiv; Rimp_. FI. Mal. Pen. 4 (1924) 347; SkotTtss. Bull. J. B. B. II, 13 (1933) 111. Outer tepals free, inner ones more or less united at the base with the filament. Anther spirally twisted. Pollengrains in tetrads. Ovary 1-locular. Capsule loculicid. Testa spirally striate. Distr. Monotypic, E.-SE. Asia, and Australia, rare in Malaysia. 1. Philydrum lanuginosum BANKS & SOL. ex GAERTN. /.c.; Mia. /.c.; HAssK. /.c.; BANKS & SOL. Bot. Coox’s Voy. 3 (1905) t. 310; Merr. Philip. J. Sc. 10 (1915) Bot. 88; Ripv. /.c.; Skottss. l.c.; YAMAMOTO, J. Soc. Trop. Agric. 10 (1938) 119; STEEN. J. Arn. Arb. 28 (1947) 420.—Fig. 1. Perennial caespitose herb, caudex short. Leaves densely rosulate, isolateral monofacial, glabrous, thick and of soft texture, 40-80 cm long incl. the sheath; sheath 14-30 by 1-1!'/2cm and 24 mm thick. Scape 1 m high or more, slender, terete, gla- brate below, villous towards the woolly inflor., with few cauline leaves gradually passing into the alternate bracts. Inflor. a terminal spike, simple or paniculate. Bracts ovate, clasping, abruptly acu- minate and subulate, 2—7 by 3/s—1 cm enclosing the buds, reflexed in anthesis, again embracing the fruit. Flowers sessile, yellow. Perianth thin, outer tepals 12-15 by up to 10 mm, acute, co-nerved, long-villous outside, margins inflexed, the posterior with 2 stronger veins and bidentate; inner petals united below 1-2 mm with the filaments, 8 by 2mm 3-nerved, spathulate, base hairy outside. Stamen 8-9 mm, glabrous; anther + spherical, 1'/2 mm across. Ovary 6—7 by 2-3 mm, densely long-wool- ly; style3-4mm, glabrous; stigma broad-triangular long-papillose. Capsule triangular-oblong; 9-10 by 45mm. Seeds co, dark-reddish, bulb-shaped, 0.8—0.9 by 0.3-0.4 mm. Distr. E. to SE. Asia (Riu Kiu Isl., Formosa, Kwantung, Hongkong, Indo-China, Siam, Burma, Andaman Isl.) and NE. Australia, in Malaysia: only in the Malay Peninsula, and in SE. New Guinea, to be expected locally elsewhere. Ecol. In ponds, marshes, and rice-fields at low altitude, in New Guinea in sedge swamps and moist savannahs. Notes. According to MERRILL (1915) the Cu- MING specimen credited to the Philippines came from the Malay Peninsula; the HILLEBRAND speci- men is certainly erroneously believed to occur wild in Java. 2. HELMHOLTZIA F. v. M. Fragm. 5 (1866) 202; Skorrss. Bot. Jahrb. 65 (1932) 260; Bull. J. B. B. mr 13 (1933) 112. Tepals united to form a short cupular tube, the inner connate to half their length with the filament. Anther straight. Pollen grains single. Ovary 3-locular. Berry leathery, (apparently) indehiscent. Seed with long funicle, outer testa lengthwise striate and not spirally so. Distr. 2 species, one in Australia, the other in E. Malaysia. [ser. I, vol. 4! FLORA MALESIANA Fig. 1. Philydrum lanuginosum BANKs, X 2/s (after BANKS & SOLANDER). Dec. 1948] PHILYDRACEAE (Skottsberg) 7 1. Helmholtzia novoguineensis (KRAUSE) SKOTTSB. Il.cc.; STEEN. J. Arn. Arb. 28 (1947) 419.— “?Xerotidae sp.’ TEYsM. Nat. Tijd. N.I. 37 (1877) 132-133.—‘Liliacea’ J. J. SmirH, Teysm. 12 (1902) 168, 329.—Astelia novoguineensis KRAUSE, Bot. Jahrb. 59 (1924) 559. Perennial herb. Rhizome stout, ascending to erect, woody, covered with leaf sheaths, up to 35 by 1-1'/2 cm; roots coarse, shoots flat, fan-shaped. Leaves densely rosulate, ensiform, 75—150 by 3—4!/2 cm; sheath 20-30 cm long, inside with scanty very long thin arachnoideous hairs, linear, acute, gla- brous, of firm texture, a bundle veins on each side forms aprominent costa dissolving to the apex, with short oblique transverse veins; blade monofacial arched or horizontal, with secondary upper and lower surface. Scape terminal 25-50 cm, erect, ob- tuse-angular, upwards covered with a dense light- grey wool, leafless in its lower half, thence carrying 5-10 reduced ensiform leaves or spathes passing into bifacial alternate bracts. Branches of Ist order of the panicle supported by a spathe, the largest 10-40 by 1!/2-3 cm, 2-3 lowermost with few branchlets of 2nd order 2-8 cm long. Bracts linear subulate 1-2 by !/4—!/2 cm, 1—3-nerved, base woolly convolute enclosing the bud. Flowers sessile, white, glabrous except the 2—2!/2 mm high tube. Outer tepals narrow-triangular, convolute with filiform apex, posterior one bicarinate-bicuspidate, with in- flexed margins, 9-12!/3 by 4-5 mm, anterior one 8-11 by 21/2-3!/2 mm. Inner tepals and filament adnate to the tube, small, 1-nerved, 4-5 mm long, irregularly 3-dentate, free portion 11/2-2!/2 by 3/4-1!/2 mm. Free part of the stamen 3!/4—33/4 mm; anther 2—2!/2 by 1-11!/2 mm. Ovary 2 by 1 mm den- sely grey-woolly. Style 3-sulcate, 23/4-4!/2 mm long, stigma small triangular. Berry white, slightly 3-sul- cate, 7-8 by 6 mm, pericarp tough leathery. Seeds ow, 2-2!/4 by 1/2 mm, _ cylindrical-flattened, often slightly curved, dark-brown with a transparent striate outer testa prolonged at both ends. Distr. Malaysia: Moluccas (Ambon, Boeroe, Ceram) and New Guinea, 600—1500 m. Ecol. In groups in muddy or moist, humic open spots in rain forests, and along ponds and margins of lakes. Fl. & fr. throughout the year. Notes. Closely allied to H. acorifolia F. v. M. from E. Australia, which has an almost glabrous scape, a more robust habit, a trifle smaller flowers, outer petals hairy on the back, style 5—6!/2 mm long, seeds mostly a little less than 2 mm. ANCISTROCLADACEAE (C. G. G. J. van Steenis, Buitenzorg) ANCISTROCLADUS (WALL. Cat. (1832) 1052) ex ARNoTT, Nov. Act. 18 (1836) 325; PLANCH. Ann. Sc. Nat. III, 13 (1849) 316; Scuerr. Nat. Tijd. N. I. 32 (1873) 407; BoerL. Handl. 1 (1890) p. XVII, XX; Kina, J. As. Soc. Beng. 42, IT (1893) 137; MaAssart, Ann. J. B. B. 12 (1895) 121; Gite, in Ey & P. ed. 2, 21 (1925) 589, f. 269-70; Riper Mal. Pen. 1 (1922) 250.—Bembix Lour. Fl. Coch. (1790) 282, nom. rej., cf .MOORE, J. Bot. 65 (1927) 279.—Wormia VAHL, Skrift. Nat. Selsk. Kjébenh. 6 (1810) 104, non ROTTB.—Bigamea KOEN. ex ENDL. Gen. Pl. 1183 (1840). Scandent shrubs (often erect in youth), without resin; branches sympodial with a series of circinate woody hooks in one plane. Leaves spread, simple, entire, often rosette-crowded, cuneiform, penninervous, reticulate-veined, glabrous, both surfaces minutely pitted, each pit with a peltate small hair secreting a wax- like substance; petiole articulated, scar on the twigs often saddle-shaped; stipules absent. Flowers 3, actinomorphic small. Inflor. few or several times dichotomous or spike-like, often provided with said hooks and single reduced bract-like leaves, branches often recurved. Pedicels articulated. Bracts with a glandular-thickened base, margin fimbriate-membranous. Calyx tube short, at length adnate to the base of the ovary; lobes 5 inequal imbricate, enlarged and wing-like in fruit. Petals 5, united at the base, slightly contorted in bud. Stamens mostly 10, rarely 5, the episepalous slightly longer. Filaments with broadened base; anthers basifixed, + introrse to + latrorse, 2-celled, opening lengthwise. Ovary for the greater part inferior, consisting of 3 carpels, 1-celled, protruding into a nipple- shaped elongation bearing 3 articulated erect styles with a punctiform or horse- shoe-shaped stigmatic apex; nipple enlarging in fruit. Ovule 1, basal, ascending, with 2 integuments. Nut not dehiscent, crowned by the enlarged calyx. Seed round- ish with testa intruding between the cerebral-like folds of the endosperm. Exocarp leathery. Embryo straight, erect, obliquely placed; cotyledons diverging; hypo- cotyl rather thick. Distr. Disjunct, ca 3 spp. in trop. W. Africa, and 9 in SE. Asia, from the Deccan to Burma, Indo- china, Hainan, S. China, the Malay Peninsula, Borneo and Sumatra (cf. fig. 2). Uses. Except for some local information nothing is known (cf. BURKILL). Ecol. In mixed rain forests, but most common on silicious soil in so-called ‘padang-scrub’, from the lowland to the hills. Kerr noted of A. wallichii (his no 7006) that all specimens grew erect, and it is reported by GAGNEPAIN to be erect in youth. R1pLey also found it on the ground as a bush, or ascending trees, and this is also observed in specimens from Sumatra and Borneo. In the open padang-scrub it is zither erect or trailing. Notes. This monogeneric family has been subsequently been referred to several families; it is now mostly placed next to the Dipterocarpaceae but differs by the 1-celled ovary, basal ovule, peculiar en- dosperm, climbing habit, sympodial structure, absence of stipules, and presence of hooks. HALLIER /. brought it to the Linaceae-Hugoniaceae, suggested already by MIQUEL. The bark of the twigs shows a peculiar cracking viz lengthwise superficial splitting of the thin grey corky outer bark and further by deeper transverse cracks. In A. extensus I found peculiar rather large crateriform glands on the base of the bracts of the inflor. Similar glands I found on 2-3 or all 5 sepals, distinctly elevated, 1-3 together. I have not found any stipules, neither in A. extensus nor in abundant living material of A. hamatus (VAHL)GILG; there are rather large bracts leaving scars amidst the leaf-tufts but these belong apparently to the leaf-spiral. GAGNEPAIN (FI. Gén. I. C. 1 (1910) 393) mentions 3-5 styles, but I found only 3. HUTCHINSON (Fam. FI. Pl. 1 (1926) 178) apparently assumes the style to be represented by the nipple- shaped extension of the ovary above the calyx on tip of which 3 free stigmas are articulated, but the tip of the latter I found distinctly ‘stigmatic papillose’ so that I assume the styles to be articulated with the ovary. The stigmatic surface is punctiform or horse-shoe-shaped. The nipple enlarges in fruit and forms a distinct part of it. All authors assume the presence of a ruminate endosperm, but HUTCHINSON§ Dec. 1948] ANCISTROCLADACEAE (Vv. Steenis) 9 denies its presence and assumes the embryo to be constituted of remarkably ‘folded cotyledons’. I had no seedlings at my disposal but an examination of the seeds did not confirm HUTCHINSON’s statement. The embryo is lying loose in the endosperm. The flowers are mostly deficient or absent in our rather rich material and when drying shrink to poor and brittle remnants. However, in A. extensus I found laterally slit anthers and not introrse cells, con- trary to GAGNEPAIN’s statements. BOERLAGE mentions slits which are turned somewhat towards the inner surface. The size of the leaves varies much both in shape and dimensions in one specimen, specially between sterile and fertile twigs. In cultivated A. hamatus I found leaves of flowering twigs 6-9 by 2—2!/2 cm, and those of sterile twigs 35-40 by 4!/2-5'/2 cm. Notwithstanding the scanty flowering material I am perfectly satisfied that only one species occurs in Malaysia. 1. Ancistrocladus tectorius (LouR.) MERR. Lingn. Se. J. 6 (1930) 329; Comm. Lour. (1935) 275.— Bembix tectoria Lour. Fl. Coch. (1790) 282.—A. extensus (WALL. Cat. 1052, nomen) PLANCH. Ann. SEs reate 4 ae = 25 = Fig. 1. Ancistrocladus tectorius (LOUR.) MERR., from Borneo, 2/s. Sc. Nat. IIT, 13 (1849) 318; Kina, J. As. Soc. Beng. 42, If (1893) 137; Boer-. Cat. pl. phan. Hort. Bot. Bog. pt 2 (1901) 114; Burk. Dict. (1935) 155.—A. pinangianus (WALL. Cat. 1054, nomen) PLANCH. lc.; Mig. Fl. Ind. Bat. 1, 2 (1859) 587; ScHEFF. Nat. Tijd. N.I. 31 (1870) 348; 32 (1873) 407; Dyer, in Hook. f. Fl. Br. Ind. 1 (1874) 300; RipL. FI. Mal. Pen. 1 (1922) 251, f. 25.—A. extensus var. pinangianus KING, J. As. Soc. Beng. 42, II (1893) 137; GaGn. l.c.; CratB, 1c.—A. hainanensis HAYATA. Ic. Pl. Form. 3 (1913) 46.—Kis. 1—2. Liana, in the youth and in open scrub often a shrub, later often trailing; main shoots provided with scattered -+ erect small leaves, between and near which arise spreading non-foliate tendril-like shoots provided with 3-6 curved hooks, lower 2 rarely 3 hooks getting woody, hooks mostly uni- lateral, rarely 1—2 alternate; these ‘tendrils’ later woody, becoming branches, upper part vanishing. Leaves crowded mostly immediately above the 2nd hook, variable in size and shape, sessile, mostly obovate-oblong, tapering towards the base, apex obtuse, rounded, acute or even acuminate, blade 9-30 by 3-10 cm; nerves 4~8 on either side, spread- ing, connected by a slightly looped intramarginal vein and a 2nd feebler outer one, rather straight, numerous secondary veins often becoming as strong as the main nerves and parallel. Inflor. be- tween the crowded leaves, very rarely lateral in the place of a ‘tendril’ on the main shoot, repeatedly dichotomous, branches divaricate, 8-15 cm long. Flowers rather crowded at their tips. Calyx lobes inequal, oval, thin-margined, glabrous except the Fig. 2. Localities of Ancistrocladus tectorius MERR. short ciliate rounded apex, some or all lobes pro- vided with 1-3 conspicuous crateriform prominent glands, mostly shorter than the corolla, 13/4—2!/2mm long, soon enlarging. Petals oblique-oval, one mar- gin often involute, acute, 3—3!/2 by 13/4 mm. Styles erect, nearly as long as the nipple-shaped ovary- 10 FLORA MALESIANA [ser. I, vol. 41, Dec. 1948] top, both 1/2 mm high, stigma punctiform. Stamens alternately inequal; filament broadened at the base; cells free, acute, more or less latrorse. Fruit with spreading calyx wings slightly decurrent on the obconical sub-5-angular smooth tube, oblong-cune- ate to spathulate, inequal, often oblique, apex blunt to rounded, with 3 larger nerves and numerous smaller densely reticulate ones, overlapping at the base, smallest mature ones measured 2!/2 by !/2cm, largest 5 by 13/4 cm; nipple broad-obcampanulate, + 3 mm high protruding, solid, not filled with part of the seed. Seed obconical with flat apex, ca 5mm high, mostly consisting of a ruminate endosperm; germ ca 2—2!/2mm high, erect, straight, obliquely inserted. ‘Distr. Burma, Siam, the Andamans, and Indo- china to S. China and Hainan, in Malaysia: Malay Peninsula, Riouw & Lingga Arch., Anambas Isl., W. Dutch Borneo, Karimata, Banka, Billiton, once collected in S. Sumatra (fig. 2). Ecol. Low altitude, often near the sea, some- times on the margin of the beach, mostly on sili- cious soils, both in mixed forest and padang scrub fr. fl. March-Aug. Vern. akar (be)boeloes, beloeloes, meloeloes (Banka), mendjoeloeng (Lepar), troeng boeloes (Billiton). Notes. I agree with BURKILL that no differences of importance can be found between A. extensus and A. pinangianus. 1 have tentatively accepted MERRILL’s name, though Moore stated that the type in the Br. Mus. is inadequate for specific iden- tification. It was collected in the classical locality but I am not satisfied that no other species grows there; in tropical regions the identification ‘by exclusion’ is a somewhat dangerous procedure. Excluded Ancistrocladus pentagynus Wars. Bot. Jahrb. 13 (1891) 385 = Durandea (Linac.) acc. to HALLIER f. (B.B.C. 39, II (1921) 68-78). Ancistrocladus sagittatus WALL. = glabra Mia. (Theac.). Tetramerista APONOGETONACEAE (C. G. G. J. van Steenis, Buitenzorg) 1. APONOGETON LinnE f. Suppl. (1781) 32; ENGL. & Krause, Pfl. R. 24 (1906). Perennial lactiferous freshwater herbs, rhizome short tuberous with fibrous roots. Leaves radical, submerged or floating, base sheathing, oblong to linear, entire or crisped, often long-petiolate; nerves lengthwise parallel, connected by numerous oblique transverse veins. Spike emerging from the water, simple or 2-8-forked, without bracts, subtended by a mostly caducous basal sheath (spathe). Flowers bisexual (rarely by abortion unisexual), small, spicate-scapose, white, rose, purple, yellow or yellowish-green. Perianth segments 2 (1-3, or absent), equal or unequal, usually persistent. Stamens in 2 rows, 6 (or more), free, hypogynous, persistent; filament filiform; anthers extrorse, small, 2-celled. Pollen subglobose or ellipsoid. Gynaecium superior, apocarpous; carpels 3-6, sessile, each with a simple style. Ovules 1-8 (or more), anatropous. Mature carpels inflated, opening along iB assice the back. Seeds without endosperm; ee outer testa often loose; embryo straight, elongate. Distr. About 40 spp. described, Africa, Ma- dagascar, Ceylon, SE. Asia, through Malaysia (very rare) to N. Australia, centering in Africa and Madagascar. Ecol. The few Malaysian specimens were col- lected in lowland stony streams both on calcareous and other rock. The testa contains in some spp. air between the two coats and float on the water; it soon decays and the embryo sinks to the bottom. Uses. The starchy tuberous rootstock is said to be edible in some spp. Notes. Monogeneric family. Next to the single indigenous species, A. fenestralis with its unique fenestral-leaved foliage is cultivated in the Bot. Gard. Buitenzorg, and may be found in private gardens as a curiosity. SS SS W993 9 S TT Eo. NTN Sas SSSaswo9 > SS Ores See ig SSS SSS S RIAA AN 1. Aponogeton loriae MARTELLI, Nuovo Giorn. Bot. Ital. I, 3 (1897) 472, t. 8; ENGL. & KRAUSE, Pfi. R. 24 (1906) 12; Domin, Bibl. Bot. 20 (1915) 254; Camus, Bull. Soc. Bot. Fr. 70 (1923) 672-3; RENDLE, J. Bot. (1923) Suppl. 58; STEEN. Journ. Arn. Arb. 28 (1947) 419.—A. crispus (non THUNB.) F. v. M. Descr. Not. Pap. Pl. 8 (1886) 51; RiDL. J. Bot. 24 (1886) 359.—A. monostachyum (non L. f.) HEMSL. Kew Bull. (1899) 113.—Fig. 1. Submerged; rootstock roundish !/2—11/2cm. Leaves green or brown, distinctly petiolate (2-15 cm), blade linear-spathulate, 10-35 by 14cm, mostly gradually tapering into the petiole, base narrow-cuneate, apex rather broadly cuneate and + blunt, primary nerves 2 on both sides and a marginal vein; paren- =SS SS aIk\ i Ai x HY, chyma opaque dotted brown-punctate; margin an "\ aa Pak \ slightly undulate-crisped to + flat. Scape 5-40 cm. yD wy FRAY Spathe ‘/2-11/2 cm long, ovate-acute, lengthwise 1 nerved, persistent, decaying gradually from the apex towards the base, green, concave, subam- Fig. 1. Aponogeton loriae MARTELLI. Plant X 1/2, plexicaulous, apex mucronulate. Flowers greenish- flower, tepal with stamen, stamen, and fruit yellow, the lower ones over 2—3 cm densely set and enlarged (after MARTELLI). iL? FLORA MALESIANA (ser. 1, vol. 4', Dec. 1948] with developing fruits, stamens about equal- ling the tepals on 2 mm long filaments, the upper ones rather abruptly as it seems male with 3 mm long stamens, very small ovaries and flowers set laxly to remote. Spikes 11/2-7 elongating in flower up to 18-20 cm. Tepals obovate ca 11/2-13/4 by 1-1/4 mm, concave, apex broadly rounded. Stamens 6; anthers roundish oval, no dehisced ones observed by me. Carpels 3, ca 2'/2 mm long, ovate, + bluntly trigonous, rather abruptly beaked by a distinct recurved rostrate style about !/2mm long. Seeds (in TEYSMANN 12792) 1-6 with a delicate loose outer coat 6-winged or -ribbed, transparently brown- reticulate-netted-celled, 2 by 2/3 mm. Inner testal coat oblong, opaque, darkbrown, smooth, 1'/4 by 1/2 mm, closely enveloping the straight embryo, easily splitting on slight pressure, rounded at both sides. Distr. Queensland (Diets 8397, n.v.), in Ma- laysia: New Guinea, and SW. Celebes in the cal- careous Maros-Pangkadjene distr. (TEYSMANN 11901, 12792). Ecol. In shallow stony streams in forests and savannahs, 100-600 m. Notes. There is a remarkable yet unexplained dimorphy in the flowers of the spike, the lower ones setting fruit only and differing in length of anthers. BURMANNIACEAE (F. P. Jonker, Utrecht) Annual or perennial, saprophytic or autotrophic herbs; the saprophytic species often colourless. Leaves usually spread or alternate, entire, simple, without stipules ; non-saprophytic species with a radical rosette of linear leaves; stem leaves often reduced to small scales; sometimes the basal part of the stem provided with many decurrent, grass-like leaves. Flowers ¢, usually actinomorphic, solitary or in capi- tate or cymose inflorescences. Perianth corolline; limb consisting of 2 whorls; tube sometimes 3-winged. Anthers 3, subsessile in the perianth throat and dehiscing laterally with horizontal slits,or 6, hanging down in the perianth tube and dehiscing with longitudinal slits. Connective large, often appendiculate. Style filiform or shortly cylindrical or conical. Stigmas 3, sometimes connate. Ovary inferior, 1-celled with parietal placentation, or 3-celled with axile placentation. Ovules «, anatropous, with 2 integuments; funicles often rather long. Fruit usually capsular, sometimes fleshy, crowned by the persistent perianth tube and the style, or by a thickened persistent basal ring of the perianth tube, dehiscing irregularly or with transverse slits at the top. Seeds ~, small, subglobose to linear, sometimes with loose, reticulate testa, with endosperm. Distr. About 125 species, widely distributed in the tropics of both hemispheres, also in subtropical America, Chicago area, Mocambique, Southern China, Japan, Southern Australia, New Zealand and Tasmania. As many species are rare, it is possible that only a part of their area is known. Most of them are found in moist regions. Among the autotrophic Malaysian Burmanniaceae there are 3 rather common species which are widely spread, viz Burmannia coelestis, B. disticha and B. longifolia. The latter two are absent from Java and the Lesser Sunda Islands, the former occurs in Java proper only in its western part. Of the saprophytic Malaysian species only 3 have been often collected, viz Burmannia championii, B. lutescens, and Gymnosiphon affinis. Ecol. The autotrophic species provided with green leaves occur in grass-fields, along road sides and river-banks, among brush-wood and in forests or on moist swampy soil, up to about 3000 m alt. The saprophytic species usually occur in dense primary or secondary forests on soils rich in humic matter by decaying wood and leaves, up to ca 1500 m alt. They are also found sometimes in bamboo bushes and parks. Notes. Treatment mostly after JONKER, A monograph of the Burmanniaceae, Thesis, Utrecht, 1938; also in: Meded. Bot. Mus. & Herb. Utrecht no 51; slightly revised. In collecting Burmanniaceae it is necessary to collect plants with complete flowers, as the limb with the stigmas and stamens is often caducous. The fruits are also important. The colour of the flowers, stems and leaves must be noted. Preservation of collections in 60 %/o spirits is recommended. In the field the saprophytic species are often found in colonies together with other saprophytic plants belonging to the Orchidaceae, Triuridaceae, and Gentianaceae. From the extreme rarity of a number of species it may be assumed that by further collecting these tiny plants several novelties will be found. KEY TO THE GENERA 1. Perianth tube cylindrical or trigonous, persistent on the capsule. Style of equal length as the tube. Anthers 3, subsessile in the perianth throat. Thecae dehiscing laterally with transverse slits TriBeE Burmannieae MIERS 2. Ovary and capsule 3-celled with axile placentation. Perianth as a whole persistent on the capsule. Ovary and perianth often prominently 3-winged, sometimes 3-costate or wingless. Ovary without glands. Capsule mostly dehiscing irregularly . . AP So 1. Burmannia 2. Ovary and capsule 1-celled with parietal placentation. Perianth limb with the stamens and stigmas deciduous. Ovary and perianth wingless. Both sides of the top of each placenta inside the ovary provided with a gland. Capsule reticulate-perforated . . . . 2. Gymnosiphon 1. Perianth tube urceolate, circumscissile, only a small basal ring persistent « on the fruit. Style very short, cylindrical or conical. Anthers 6, hanging down in the tube. Thecae dehiscing introrsely with longi- tuidinalishits =< = . Stace TriBE Thismieae MIERS 3. Inner perianth lobes free, « or converging at their tops « or connate to a mitre with 3 holes, the latter without appendages at the apex ; ~ « 1 e See hismia 3. Inner perianth lobes connate to a mitre with 3 holes, crowned by 1 or 2 appendages. 4. Mitre crowned by 3 erect, thick, filiform appendages, clavately swollen at their tops . 4. Geomitra 4. Mitre crowned by 1 erect thick column, bearing at its apex 3, more or less connate, glandular MODES wher Os ae ee Tae SA ake a a en een. eae ee See SCH Nii DOE 14 FLORA MALESIANA [ser. I, vol. 4! Fig. 1-8. Burmanniaceae. 1. Burmannia bifaria J. J. S., xX 4, 2. Scaphiophora gigantea JONK., X 1/3, 3. Thismia aseroe BeEcc., X 8/s, 4. Th. episcopalis (BEcc.) F. v. M., x 2/3, 5. Burmannia coelestis Don, 3/2, 6. Burmannia championii Tuw., X 1/1, 7. Gymnosiphon aphyllus Bu., < 2/3, 8. Burmannia longifolia BEcc., x 2/s. Dec. 1948] BURMANNIACEAE (Jonker) i 15 1. BURMANNIA LINNE, Sp. PI. ed. 1 (1753) 287; JONKER, Monogr. (1938) 18, 57. Annual or perennial, saprophytic and colourless or chlorophyllose. Flowers often 3-winged. Perianth limb usually consisting of 6 lobes; the outer ones being much larger; inner 3 often minute, sometimes lacking. Perianth tube cylindrical to trigonous. Anthers 3; connective sometimes with 2 apical crests and/or a han ging, median, basal spur. Style filiform, branching into 3 short branches, each bearing a stigma, or 3 sessile stigmas at the apex of the style. Ovary trigonous. Fruit cap- sular, mostly dehiscing irregularly. Seeds many, oblong or ellipsoid. Distr. 57 species, tropics of both hemispheres, also in the S. United States, S. part of S. America Mocambique, S. China, Japan and S. Australia. KEY TO THE SPECIES 1. Perennial, leafy green herbs. Greater part of the stem beset with grass-like, linear or ensiform, de- current, imbricate leaves. Inflorescence usually many-flowered. Flowers hanging, very narrowly a mecmiatmehe Basal part) ss «2M. 2 es (Sect. Foliosa Jonx.). 1. B. longifolia 1. Annual or perennial, saprophytic or green herbs. Stem leaves reduced to small scales. The non- saprophytic species with a radical rosette of linear leaves. Stem usually 1- to few-flowered. Flowers Peres fae is, rah ee ew OOS. Mel Son 8 QSEeR Enburmannia MALMeE) 2. Non-saprophytic, chlorophyllose herbs with a rosette of green leaves at the base; rosette often consisting of only 1-3 leaves. 3. Flower wings narrower than the perianth tube or reduced to ribs. 4. Basal rosulate leaves few. Stem bearing 1-2 flowers at its apex. Connective with 2 apical crests, basal spur lacking. Ovary as long as the perianth or longer . . . . . 2. B. geelvinkiana 4. Basal rosette well developed. Stem bearing at its apex a usually bifid inflorescence. Connective provided with 2 apical crests and a basal, hanging spur. Ovary shorter than the perianth 3. B. bancana 3. Flower wings as wide as the perianth tube or wider. 5. Margin of the perianth lobes double. Connective with a basal hanging spur and 2 apical crests. Thecae separated. 6. Robust herbs with a well developed rosette of grass-like, up to 15 cm long leaves. Inflorescence msually a bifid, many-flowered'cyme 95. ss ee B. disticha 6. Slender herbs. Basal rosulate leaves about 1 cm. Stem bearing at its apex a single flower or a PumeMRICM TIO CIS TT OUIH) ili eke Pe ht a ee 5. B. coelestis 5. Perianth lobes with single margin. Connective with 2 apical crests; basal spur lacking. Thecae connate below the basal connective margin . ........ ., . 6. B. connata 2. Saprophytic herbs without chlorophyll. Radical rosette absent. 7. Flowers wingless, 3- or 6-costate. 8. Stem scales many, imbricate in the lower part of the stem. Ovary as long as the perianth or JNpoSh 5 0S Sa eer mere) es rae ree 1 em rN 0p rs Se . . 7. B. sphagnoides 8. Stem scales not imbricate. Ovary shorter than the perianth. 9. Flowers 6-costate. Perianth limb thick, fleshy, more or less succulent ft £534 8. B. bifaria 9. Flowers 3-costate to narrowly 3-winged. Limb not fleshy. 10. Inflorescence usually capitate. Inner perianth lobes spathulate, sometimes slightly papillose. Connective mucronate at the apex, obtuse at the base . . - « = 49. B. championii 10. Flowers usually pedicellate. Inner perianth lobes broadly obovate, distinctly papillose. Con- nective not mucronate, acute atthe base . . . .. . Sede A 10. B. micropetala 7. Flowers 3-winged. 11. Inner perianth lobes absent. 12. Perianth lobes simple. Connective with an apical, papillose crest and a basal, hanging, obtuse spur ee te he Meh aloe ae Mi OR ls. oe ms Ue ABA Libs ae oT ia tridentate 12. Perianth lobes bifid. Connective without crest and Spurte alee werkt 12. B. oblonga 11 Inner perianth lobes not lacking, sometimes very small. 13. Very delicate plants. Connective with a basal) hansingispure-.. 2005 . 13. B. steenisii 13. Plants not very delicate. Connective without basal, hanging spur. 14. Stem rather robust, often many-flowered. Inner perianth lobes minute, orbicular. Flower wings variable, linear to half cuneate or quadrangular . . . . . . 14. B. lutescens 14. Slender herbs, usually 1-flowered. Inner perianth lobes lanceolate. Flower wings elliptical 15. B. malasica 16 FLORA MALESIANA [ser. I, vol. 4! 1. Burmannia longifolia Becc. Malesia 1 (1877) 244; JoNKER, Monogr. (1938) 20, 59.—B. leucantha Scuutr. Bot. Jahrb. 49 (1913) 107.—Fig. 8. Perennial, 12-50 cm. Stem usually simple, forked at the top into the bifid inflorescence or bearing a simple cincinnus. Leaves linear, sometimes keeled, decurrent, stem-clasping, acute, sometimes subul- ate, parallel-veined but midrib more prominent, growing smaller towards the top, 4-20 cm by 2-9 mm. Upper part of stem beset with appressed, scattered, lanceolate, acute scales, 5-33 mm. Basal part of stem with brownish, dried leaves. Inflo- rescence 32-1-flowered, branches up to 4cm. Bracts. scale-like, linear-lanceolate, 5-10 mm. Flowers subsessile, hanging, white, often with pale- violet or bluish limb, 8-16 mm. Outer perianth lobes deltoid, acute, 2-4!/2 mm; margin fleshy at the base. Inner ones broad-obovate to orbiculate, entire and rounded, or retuse, or bilobate, 1!/2-2 mm. Perianth tube cylindrical, sometimes swollen in the upper part, 3-5 mm; lower part of tube and ovary very narrowly 3-winged. Stamens inserted just be- low the inner perianth lobes. Connective broad, oblong, crowned by two, rather wide crests. Fila- ments short, broad. Style thick, branching into 3 very short branches, each bearing a curved funnel- shaped stigma. Ovary obovoid, 4-7 mm. Capsule obovoid, dehiscing transversally, irregularly. Seeds oblong to scobiform, appendaged at both sides; testa loose, reticulate. Distr. All over Malaysia, except Java and the Lesser Sunda Islands. Ecol. In mountain forests, brush-wood, along mossy trails, often on ridges, scattered, ascending to 2800 m alt. 2. Burmannia geelvinkiana Becc. Malesia 1 (1877) 244: JoNKER, Monogr. (1938) 111. Annual, 7-12 cm. Stem filiform, simple, bearing 1 or, sometimes, 2 flowers. Rosulate leaves 2-5, linear, subulate, 3-nerved, 3-5 mm by 1 mm. Stem leaves scale-like, appressed, linear-lanceolate, acuminate to subulate, up to 3 mm. Bracts ovate- lanceolate, long-acuminate, 3-nerved, 1!/2 mm. Flowers bluish, very narrowly 3-winged, 7 mm. Outer perianth lobes triangular to broad-ovate, apiculate, about 11/2 mm. Inner ones linear, obtuse, 1/2-1 mm. Connective thick, triangular, obtuse at the base, bearing two divergent, slightly papillose crests at the apex. Style rather short and thick, bearing 3 sessile, funnel-shaped stigmas; style with stigmas about 1!/2 mm. Ovary ellipsoid to narrowly obovoid, about 4 mm. Flower wings linear, about 51/2 mm by !/4 mm. Capsule obovoid, dehiscing with transverse slits. Seeds ovoid, bright yellow. Distr. Malaysia: West New Guinea (Wandam- men Peninsula, Geelvink Bay), once collected. 3. Burmannia bancana Mia. FI. Ind. Bat. Suppl. 1 (1860) 617; JonKER, Monogr. (1938) 24, 113.—B. graminifolia WARB. in FEDDE, Rep. 18 (1922) 330. Annual, 20-37 cm. Stem simple, terete, forked into the inflorescence. Rosette distinct; /eaves many, linear to lanceolate, subulate, parallel-veined, 2!/2-7 cm by 6 mm. Stem leaves few, scale-like, ap- pressed, lanceolate, subulate, 1-3 cm. One rosette sometimes bearing 2 or 3 stems. Inflorescence a double cincinnus, 3- to 5-flowered; branches up to 3 cm. Flowers blue or purplish, narrowly winged, 6— 13 mm. Outer perianth lobes lanceolate-triangular, — acute, with 3 prominent, fleshy nerves inside, up — to 3 mm. Inner ones linear-lanceolate, obtuse, up © to 2'/2mm; midrib prominent, fleshy. Perianth © tube cylindrical-trigonous, up to 41/2 mm. Connect- ive oblong, provided with a basal, hanging, obtuse © spur and 2 apical, divergent obtuse crests. Style — filiform, branching into 3 short branches, each © bearing a slightly curved, funnel-shaped stigma. Ovary truncate-ellipsoid, 3-7 mm. Flower wings linear, 11 by ‘'/2-1 mm. Capsule ellipsoid to obovoid. Testa of the seeds elongate. Distr. Malaysia: Sumatra, Banka, Billiton, Borneo. Ecol. Wet places, along streams, &c. Vern. Roempoet taroem, oemboet oemboet (Bil- liton). 4. Burmannia disticha LINNE, Spec. Plant. 1 (1753) 287; JoNKER, Monogr. (1938) 115.—B. distachya R. Br. Prod. Fl. Nov. Holl. 1 (1810) 265.—B. suma- trana Mia. FI. Ind. Bat. Suppl. 1 (1860) 616.—B. disticha var. sumatrana Hook. f. Fl. Br. Ind. 5 (1888) 664. Robust annual, up to 75 cm. Stem usually simple, forked into the inflorescence. Rosette distinct; leaves linear or lanceolate, acute, up to 15 cm by 13 mm. Stem leaves reduced to appressed, lance- olate, acute or acuminate scales, up to 7cm by 7 mm, imbricate in the lower part of stem; upper part of stem often leafless. Inflorescence branches up to 8cm. Bracts lanceolate, acute, about 5-12mm. Flowers sessile or shortly pedicellate, blue or pur- plish, often with yellow-tipped, greenish lobes, rarely yellow, 10-20 mm. Outer perianth lobes tri- angular, acute, 2'/2mm; margin thick, double in the basal part. Inner lobes linear-lanceolate, fleshy, obtuse, 1—-1!/2 mm. Perianth tube cylindrical-trigo- nous, 3—4!/2 mm. Connective broad, provided with 2 distinct, acute apical crests and a broad, obtuse to almost truncate, basal, hanging spur. Style thick- filiform, bearing 3 sessile, funnel-shaped stigmas; style with stigmas about 3 mm. Ovary ellipsoid to obovoid, truncate, attenuate towards the base, up to 1 cm. Flower wings elliptical, 10-18 by 1!/2-21/2 mm, continuing as crests on the back of the outer perianth lobes, decurrent along the short pedicel. Capsule obovoid, truncate, irregularly dehiscing with transverse slits. Distr. Widely distributed in the tropics of Asia and Australia: Ceylon, India, Siam, Indo-China, China, through Malaysia to Australia, in Malaysia hitherto not found in Java, the Lesser Sunda Is- lands, Moluccas, and Philippines. Ecol. A species with a large ecological ampli- tude. It has been collected in brush-wood, swamps and bogs among Sphagna, moist hollows, open grasslands, mountain meadows, marshy plateaus, on bare rocks, and has even been recorded as growing in water; ascending to ca 3500 m alt. Vern. Si goeroe goeroe (Sumatra). Dec. 1948] BURMANNIACEAE (Jonker) . 17 5. Burmannia coelestis Don, Prod. Fl. Nep. (1825) 44; JonKER, Monogr. (1938) 120.—B. javanica BL. Enum. FI. Jav. 1 (1827) 28.—B. triflora Roxs. FI. Ind. 2 (1832) 117.—B. azurea GrirF. Not. 3 (1851) 326.—B. selebica Becc. Malesia 1 (1877) 243.—B. borneensis, B. chinensis, B. malaccensis & B. rigida GANDOG. Bull. Soc. Bot. Fr. 66 (1919) 290.—Fig. 5. Autotrophic annual, up to 30 cm. Stem simple or, sometimes, branched, bearing a single flower or a cluster of few flowers. Rosulate /eaves linear or lanceolate, acute or acuminate, 3-nerved, about 1 cmby 1!/2-3 mm. Stem leaves appressed, imbricate in the basal part, linear-lanceolate, subulate, rather long, up to 2cm. Bracts lanceolate, acute, 4 by 1 mm. Flowers prominently 3-winged, blue, pur- plish or white, often with yellow lobes, about 11'/2 mm. Outer perianth lobes ovate, apiculate, with double margin, about 1!/2 mm. Inner ones lanceolate, apiculate, with double margin, about 1/2 mm. Tube cylindrical-trigonous, about 5 mm. Connective provided with 2 apical, divergent, ob- tuse to truncate crests and a basal hanging, rather long, obtuse spur. Style thick-filiform, bearing 3 sessile, funnel-shaped stigmas with swollen margin. Style with stigmas about 4 mm. Ovary ellipsoid to obovoid, truncate, attenuate towards the base, about 5mm. Flower wings half elliptical to half obovate, about 10 by 2!/2 mm. Capsule obovoid, truncate, transversely dehiscing. Distr. Widely spread in tropical Asia: India, Siam, Indo-China, S. China, and the Caroline Is- lands, in Malaysia throughout the Archipelago, in Java only in the W. part, once collected in the S. part of Madoera Island, and once in Bali, other- wise absent from the Lesser Sunda Islands. Ecol. Grass-fields, among alang-alang (Jmpe- rata), in mountain meadows, parks and plantations, ascending to ca 1700 m alt. 6. Burmannia connata JONKER, Monogr. (1938) 128. Autotrophic annual, 15-30 cm. Stem simple, bearing 1-3, shortly pedicellate flowers. Rosulate leaves few, linear, acute or acuminate, 1-nerved, 4-8 by 1mm. Stem leaves scale-like, appressed, linear, acute, 2-S mm. Bracts linear-lanceolate, acute, 1'/2 mm. Flowers prominently 3-winged, 6-8 mm. Outer perianth lobes acute, with involute margin, 1 mm. Inner ones broadly ovate, obtuse, nearly 1/2 mmlong. Perianth tube cylindrical, 3 mm. Connective rather broad, provided with 2 apical, divergent, obtuse crests. Thecae bright yellow, ap- pressed against the connective and connate below the basal connective margin. Basal hanging spur lacking. Style as long as the tube, branching into 3 short branches, each bearing a peltate, disk- shaped stigma. Ovary ellipsoid to obconical, 2'/2-4 mm. Flower wings half oblanceolate, decur- rent along the pedicel, 8 by 2mm. Capsule ellipsoid, dehiscing with transverse slits ,4—6 mm. Seeds scobi- form, yellow. Distr. Malaysia: Sumatra, Eastcoast Resi- dency (Kota Pinang, Soengei Kana). 7. Burmannia sphagnoides Becc. Malesia 1 (1877) 246; JONKER, Monogr. (1938) 135. Saprophyte, 4!/2-12cm. Stem simple, thick, beset with many lanceolate, acute, in the lower part im- bricate, 3-6 mm long, scale-like /eaves and bearing 2-5 subsessile flowers at the apex. Bracts broad- lanceolate to ovate, acute, 3-6 mm. Flowers 6-cost- ate, white, about 8!/2mm. Outer perianth lobes broadly triangular, with swollen margin, about 8!/2 mm. Inner ones fleshy, obtuse to rounded, papillose, almost 1 mm. Perianth tube cylindrical, about 2 mm. Connective oblong, acute at the base crowned by 2 divergent, obtuse crests. Style thick- filiform, bearing 3 sessile, obconical stigmas. Ovary large, broadly ellipsoid to subglobose, 4-5 mm. Distr. Malaysia: Malay Peninsula, Sumatra (Eastcoast Res.), and Borneo (Sarawak). Ecol. A rare species, occurring in decaying matter in forests. 8. Burmannia bifaria J.J.S. Icon. Bogor. 4 (1914) 379; JONKER, Monogr. (1938) 136.—B. engganensis JONKER, Blumea 3 (1938) 108; Monogr. (1938) 137. —Fig. 1. Saprophyte, 5-13 cm. Stem simple or branched, beset with scale-like, ovate to lanceolate, 1-nerved, sometimes distichous, up to 5 mm long /eaves and bearing 1 flower or branching into a bifid, up to 9-flowered cyme. Flowers shortly pedicellate, 6- costate, white or somewhat purplish, 9-13 mm. Limb fleshy. Outer perianth lobes triangular, ob- tuse, with involute, crenate margin, 1!/2-2 mm. Inner ones ovate to orbicular, !/4—1'/2 mm. Perianth tube cylindrical-trigonous, 2!/2-5 mm. Connective obtri- angular, crowned by 2 divergent, papillose crests. Style thick-filiform, branching into 3 short branch- es, each bearing a somewhat funnel-shaped stigma with a broad, rotundate, membranous, hanging appendage. Style with stigmas 4-4!/2 mm. Ovary ellipsoid, 3-6 mm. Seeds ovoid, brown. Flower wings reduced to narrow, linear ribs. Distr. Malaysia: West Java (vicinity of Buiten- zorg) and Enggano Island (off SW. Sumatra). Ecol. Among decaying leaves in forests, as- cending to 1000 m alt. 9. Burmannia championii THw. Enum. Pl. Zey!l. (1864) 325; JoNKER, Monogr. (1938) 138.—B. tuberosa Becc. Malesia 1 (1877) 245.—B. capitata (non MART.) MAKINO, Bot. Mag. Tok. 4 (1890) 23. —B. japonica MAxIM. ex Mak. Ill. Fl. Jap. 1, no 7 (1891) 4.—B. dalzieli RENDLE, Journ. Bot. 40 (1902) 311.—B. chionantha SCHLTR. Bot. Jahrb. 49 (1913) 107. Fig. 6. Saprophyte, 2-18cm. Rhizome tuberous or elongate, covered with hair-like roots, producing small, adventitious tubers. Stem simple, beset with lanceolate, acute, appressed, scale-like, 142-4 mm long leaves. Bracts lanceolate, acute, about 3 mm. Flowers subsessile in a capitate inflorescence at the top of the stem, 3-costate, white, 5-12 mm. Outer perianth lobes triangular, acute, with involute margin in the upper part, 1—2!/2 mm. Inner ones spathulate, rounded, slightly papillose at the mar- gin, about 3/4 mm. Connective broadly oblong, ob- tuse at the base, crowned by 2 indistinct, divergent, obtuse crests and provided with a median small 18 FLORA MALESIANA [ser. I, vol. 4! point at the apex, usually directed inwards and then hardly perceptible. Style thick-filiform, bearing 3 subsessile funnel-shaped stigmas; style with stig- mas 3 mm. Ovary ellipsoid to obovoid, 2-3 mm. Fig. 9. Burmannia lutescens BeEcc., with broad wings (Mt Gedeh, W. Java), x 1/1. Distr. Ceylon, S. China, Japan and Malaysia: Batoe Islands (off W. Sumatra), Banka, W. Java, Borneo, and New Guinea. Ecol. A species with a large ecological ampli- tude, occurring in humus of moist forests and also in parks, plantations, bamboo bush, on rocks in streams. 10. Burmannia micropetala RIDL. Trans. Linn. Soc. II, 9 (1916) 228; JonKER, Monogr. (1938) 140. Saprophyte, 7!/2-15 cm. Stem simple, beset with acute or acuminate, 1-nerved, often keeled, 2-5 mm long, scale-like /eaves. Bracts linear-lance- olate, acute, 1-nerved, about 4 mm. Pedicels up to 5mm. Flowers shortly pedicellate, in contracted 3—8-flowered bifid or, sometimes, pseudo-umbel- late cymes, very narrowly 3-winged to 3-costate, 7-9 mm. Outer perianth lobes triangular, acute, about 2 mm, in the upper part provided with small, rounded, crenate lateral lobes. Inner ones broadly obovate, rounded, papillose at the margin, !/2 mm. Perianth tube cylindrical, about 3 mm. Connective oblong, acute at the base, crowned by 2 divergent, acute crests. Style thick-filiform, branching at the apex into 3 very short branches, each bearing a funnel-shaped stigma with 2 small, apical points. Style with stigmas about 3 mm. Ovary ellipsoid, truncate, about 2!/2 mm. Distr. Malaysia: New Guinea only. 11. Burmannia tridentata Becc. Malesia 1 (1877) 246; JONKER, Monogr. (1938) 141. Saprophyte, 6-14cm. Stem simple or branched, beset with appressed, lanceolate, acute, 1-veined, slighly keeled, scale-like, 11/2-2 mm long leaves. Bracts ovate-lanceolate, acuminate, 1-veined, about 1!/2mm. Stem or branches bearing 1-3, pro- minently winged, 2-7 mm long flowers. Outer peri- anth lobes triangular to ovate, obtuse, swollen at the margin, 1—11/2mm. Inner lobes absent. Perianth tube cylindrical, about 2mm. Connective quad- rangular with a broad, swollen, obtuse, hanging, basal spur, and an apical, erect, papillose, obtuse crest. Style as long as the tube, bearing 3 subsessile, funnel-shaped stigmas. Ovary subglobose, about 2mm. Flower wings half elliptical to half-orbi- culate, about 4 by 2mm. Capsule subglobose, about 2!/2 mm. Seeds scobiform. Distr. Malaysia: Borneo, Sarawak (Mt Mat- tang), once collected. 12. Burmannia oblonga RIDL. J. Str. Br. R. As. Soc. 41 (1904) 33; JoNKER, Monogr. (1938) 25, 142.— B. bifida GAGNEP. Bull. Soc. Bot. Fr. 54 (1907) 462. Saprophyte, 7-15 cm. Stem simple or branched, bearing 1—2 flowers, beset with appressed, ovate to lanceolate, obtuse, scale-like, about 1!/2mm long leaves. Below the flower 2 lanceolate, scale-like bracts, 2!/2mm. Flowers white, sometimes with yellow limb, 8-10 mm. Outer perianth lobes bifid, obtuse, about 11/2 mm, papillose in the upper half at the margin, in the lower half with 2 yellow bags inside, provided with 2 involute, narrow triangular lateral lobes. Inner ones absent. Tube conical, 4-4!/2mm long. Connective oblong, yellow, without crests or spur. Style thick-filiform, bearing at the apex 3 sessile, funnel-shaped curved stigmas. Style with stigmas about 4!/2mm. Ovary subglobose, 2!/2-4 mm. Flower wings obovate, truncate, white, 5-7'/2 by 3-4 mm. Distr. Hainan, Indo-China and Malaysia: Ma- lay Peninsula, N. Sumatra (Atjeh and Eastcoast Res.). Ecol. On forested rocks or loamy soil in dense jungle or forest, ascending to 1300 m. 13. Burmannia steenisii JONKER, Monogr. (1938) 158. Delicate saprophyte, 2-6 cm. Stem simple or branched, bearing 1—2 flowers, beset with lance- olate, acute, scale-like, !/2-11/2mm long /eaves. Below each flower 1 or 2 lanceolate, 1-veined, acute bracts, about 1'/2mm. Flowers pure white with yellow Dec. 1948] BURMANNIACEAE (Jonker) 19 limb, prominently 3-winged, 3-7 mm. Outer peri- anth lobes triangular, subobtuse, with swollen mar- gin, about 1 mm. Inner ones orbiculate, minute. Perianth tube cylindrical-trigonous to conical-trig- onous, about 2!/2 mm. Connective quadrangular, with a broad, obtuse, basal hanging spur and crowned by 2 short, thick, straight, obtuse, di- vergent crests. Style thick-filiform, bearing 3 ses- sile, slightly curved, bilabiate, funnel-shaped stig- mas. Ovary subglobose, about 2 mm. Flower wings half elliptical to half-quadrangular, pure white, about 4!/2 by 1!/2mm. Capsule subglobose, dehiscing with transverse slits. Seeds scobiform. Distr. Malaysia: E. Java, Pasoeroean Resi- dency (Mt Lamongan). Ecol. Collected on the SW. slope of Mt La- mongan on coarse, volcanic sandy soil in brush- wood, ca 600 m alt. It is the only species of the family hitherto reported from East Java. 14. Burmannia lutescens Becc. Malesia 1 (1877) 246; JONKER, Monogr. (1938) 24, 148.—Gonianthes candida BLUME, Cat. Gew. Buitenzorg (1823) 20.— Gonyanthes candida BLUME, Flora 8 (1825) 123.— B. candida (Bu.) ENGL. Nat. Pfl. Fam. 2,6 (1889) 50, not B. candida Grirr. ex Hook. f.—B. gracilis Ripe. J. Sir. Br: R. As. Soc. 22 (1890) 335.—B. papillosa STAPF, Trans. Linn. Soc. II, 4 (1894) 232. —B. novae-hiberniae SCHLTR. in K. Scu. & LAuT. Nachtr. Fl. D. Sch. Geb. (1905) 73.—B. gjellerupii J.J.S. in Feppe, Rep. 10(1912) 487.—B. gonyantha Hocur. Candollea 2 (1925) 325.—Fig. 9-10. Saprophyte, up to 23 cm. Stem thickly filiform to robust, simple or branched, 1-flowered or forked into the inflorescence. Leaves lanceolate to ovate, acute, 1—3!/2 mm. Bracts lanceolate to ovate, often keeled and carinate. Cincinni bifid, up to 11-flow- ered; branches up to 3cm. Flowers pedicellate, white, sometimes with yellow limb, seldom bluish, about 8'/2mm. Outer perianth lobes ovate or tri- angular, apiculate, about 1!/2 mm; margin fleshy. Inner lobes minute, fleshy, orbiculate. Perianth tube trigonous, 2!/2-5 mm. Connective truncate, rounded at the base, slightly 2-lobed at the apex into 2 very short, papillose crests. Basal spur ab- sent. Style thick, bearing 3 subsessile, funnel- shaped to bowl-shaped stigmas. Style with stigmas as long as the tube. Ovary subglobose to truncate- globose, 3-5 mm. Flower wings various, linear, or elliptical, or rather broad, half-cuneate or qua- drangular, running from the base of the limb to the middle or the base of the ovary. Capsule subglo- bose, dehiscing with large horizontal slits. Distr. Malaysia: Sumatra, Malay Peninsula, Borneo, Java, New Guinea, and New Ireland. Ecol. Usually in the humus of shady moist forests, up to 1500 m. Notes. Specimens with narrow perianth wings have been described as B. lutescens, B. novae-hiber- niae, B. gjellerupii, with elliptical ones as Goni- anthes candida, and rather broad-winged speci- mens as B. gracilis. They all belong to one species variable in that respect. In Java the species was often called B. candida (BL.) ENGL. but this is a later homonym; B. candida Grirr. ex HOOK. f. is Fig. 10. Burmannia lutescens Becc., Mt Salak, W. Java. Form with narrow perianth wings, x 2/3. an allied species, occurring in Burma, Siam and the Langkawi Islands. 15. Burmannia malasica JoNKER, Monogr. (1938) 152.—Burmannia lutescens (non BEcc.) WINKLER, Bot. Jahrb. 48 (1913) 96. Saprophyte, 5!/2-8 cm. Stem simple, 1-flowered, seldom 2-flowered, beset with few appressed, lance- olate, subacute, 1-veined, slightly keeled, 1!/2-2 mm long, scale-like /eaves. Bracts elliptical, acuminate, l-veined, 1'/2 mm. Flowers purple or white with yellow limb, prominently winged, about 5 mm. Outer perianth lobes triangular with swollen mar- gin, acuminate to apiculate, about 1 mm. Inner lobes erect, lanceolate-ovate, obtuse, about !/2 mm. Perianth tube cylindrical, 1!/2 mm. Connective obtriangular, obtuse at the base, provided with 2 short, divergent crests at the apex. Style cylindrical, bearing 3 sessile, funnel-shaped stigmas. Ovary subglobose to ellipsoid, 2!/2by 2 mm. Flower wings half-orbiculate to half-elliptical, about 4 by 2 mm. Capsule ellipsoid to obovoid, dehiscing with a transverse slit. Seeds scobiform to fusiform. Distr. S. Siam and Malaysia: SE. Borneo. 20 FLORA MALESIANA [ser. I, vol. 4! 2. GYMNOSIPHON BLuME, Enum. PI. Jav. I (1827) 29; JONKER, Monogr. (1938) 27, 168. Saprophytic annuals, without chlorophyll. Leaves scale-like. Perianth limb con- sisting of 6 lobes, the 3 outer being much larger and slightly 3-lobed. Anthers 3, sessile in the throat. Thecae bursting horizontally. Ovary ovoid to globose, 1-celled with 3 parietal placentas, each placenta provided with a large, globose gland at both sides of the top. Style filiform, branching into 3 short branches, each bearing a stigma. Perianth limb, stamens and the upper part of style with the stig- mas caducous after flowering. Capsule crowned by the persistent perianth tube. Seeds ovoid to globose, reticulate. Distr. 29 species, tropics of both hemispheres, not in Australia. Notes. In Asia, and Malaysia, this genus is represented by the section Eugymnosiphon URBAN only, characterized by the reticulate-perforated capsule dehiscing at the top. KEY TO THE SPECIES 1. Anthers inserted above the middle of the perianth. 2. Flowers pedicellate, in loose, many-flowered cincinni or bifid cincinni 1. G. aphyllus 2. Flowers + sessile in a 1- or sparsely flowered inflorescence. 3. Capsule + globose. Margin lobes of the outer perianth lobes crenate . 3. Capsule conical-ovoid. Margin lobes of the outer perianth lobes entire 2. G. oliganthus 3. G. minahassae 1. Anthers inserted in or below the middle of the perianth. 4. Anthers inserted below the middle of the perianth. Connective apiculate at the top. Ovary elongate- conical, tapering to the pedicel 4. G. affinis 4. Anthers inserted in the middle of the perianth. Connective not apiculate. Ovary marked from the pedicel. 5. Flowers sessile or subsessile in loose cincinni or bifid cincinni . 5. G. papuanus 5. Stem 1—2-flowered, or many sessile flowers in a capitate inflorescence at the top of the stem. 6. Outer perianth lobes acuminate, a third of the length of the whole perianth. Connective narrow. Stigmas with dorsal crest. Inflorescence 2- to many-flowered 6. G. neglectus 6. Outer perianth lobes deltoid, short, a fifth of the whole peeene Stiemas without crest. Inflo- rescence 1—3-flowered 1. Gymnosiphon aphyllus BLUME, Enum. PI. Jav. 1 (1827) 29; JoNKER, Monogr. (1938) 30, 170.—G. borneense Becc. Malesia 1 (1877) 241.—G. pedicel- latum SCHLTR. Bot. Jahrb. 49 (1913) 105.—Fig. 7. Stem up to 17 cm, forked into a bifid cincinnus or bearing a simple cincinnus. Leaves acute, often keeled, appressed, 1—21/2 mm. Bracts ovate, obtuse, scale-like. Pedicels 1-5 mm. Inflorescence often loose and many-flowered. Perianth white or lilac; tubular part up to 4mm; limb about 2!/2 mm. Outer perianth lobes ovate, obtuse, provided with a narrow, crenate lateral lobe at both sides. Inner ones linear-lanceolate, minute. Stigmas curved, funnel-shaped, inappendiculate. Capsule about 3 mm. Distr. S. Siam, throughout Malaysia. Ecol. On humus or decaying wood or leaves in the shade of moist forests, below 1500 m alt. 2. Gymnosiphon oliganthus SCHLTR. Bot. Jahrb. 49 (1913) 101; JonKEeR, Monogr. (1938) 172. Stem tender, simple or branched, 4—9!/2 cm, 1- or 2-flowered. Leaves and bracts minute, to 1 mm, keeled and appressed. Pedicels up to 3 mm. Flowers white to bluish lilac, up to 5 mm. Outer perianth lobes triangular, subobtuse, provided with crenate lateral lobes. Inner ones small, cuneate, obtuse to 7. G. pauciflorus Distr. Malaysia: NE. Brit. New Guinea, once collected. Ecol. In forests, 450 m alt. 3. Gymnosiphon minahassae SCHLTR. Bot. Jahrb. 49 (1913) 104; JoNnKER, Monogr. (1938) 172. Stem usually simple, 2—5-flowered, 7-12 cm. Leaves acute, appressed, up to 1 mm. Bracts more or less obtuse, keeled. Pedicels 1-3 mm. Perianth white with bluish limb. Outer perianth lobes ob- tuse; lateral lobes entire. Inner ones lanceolate. Perianth limb deciduous above the anthers. Stigmas quadrangular, truncate at the apex, apiculate at the base. Distr. Malaysia: N.Celebes (Minahasa), once collected. Ecol. In humic soil, 800 m alt. 4. Gymnosiphon affinis J.J.S. Nova Guinea 8 (1909) 194; JonKER, Monogr. (1938) 31.—G. tor- ricellense SCHLTR. Bot. Jahrb. 49 (1913) 101. Stem 7-13 cm, simple or branched, white, lilac — or rose-coloured, forked into a bifid cincinnus or — bearing a simple cincinnus of 1-3 flowers. Leaves — ovate, acuminate, 1-2 mm. Bracts to 3 mm. Pedi- — cels 14mm. Perianth white, limb 2!/2 mm, tube — 1'/2mm. Outer perianth lobes ovate, obtuse; lateral — truncate. Stigmas auriculate, soup-plate-shaped. lobes crenate. Inner ones small, rather broad, — | Dec. 1948] BURMANNIACEAE (Jonker) oH obovate, obtuse. Tube swollen at the insertion of the stamens. Connective quadrangular, acute-api- culate at the apex. Stigmas rather large, soup-plate- shaped. Ovary obconical, swollen in the upper part, about 1'/2 mm. Capsule thick-ellipsoid, about 3-3!/2 mm, crowned by the short, cylindrical to conical, 2mm long, persistent part of the tube. Seeds brownish, fusiform, reticulate. Distr. Malaysia: New Guinea. Ecol. In forests, in humic soil, ascending to ca 700 m alt. 5. Gymnosiphon papuanus Becc. Malesia 1 (1877) 241; JoNKER, Monogr. (1938) 174.—G. celebicum ScHLTR. Bot. Jahrb. 49 (1913) 104. Stem simple or branched, 4-14 cm, colourless, bearing rather loose simple or bifid cymes of 3-many subsessile flowers. Leaves acuminate, up to 2!/2 mm. Bracts up to 3!/2 mm. Perianth whitish- purplish; limb about 1!/2 mm; tube up to 4!/2 mm. Outer perianth lobes ovate, triangular, obtuse; lateral lobes entire, involute. Inner ones small, linear. Connective deltoid, at the top 3-lobed, pro- vided with a forked thickening. Stigmas rather large, soup-plate-shaped, obtuse, cordate, auricu- late at the base. Ovary ovoid, about 1!/2 mm. Cap- sule obovoid to truncate-subglobose, 2!/2-5 mm long; crowned by the 2-2!/2 mm long, cylindrical persistent part of the tube. Distr. Micronesia (Palau Islands), in Malaysia: Celebes and New Guinea. Ecol. Moist forests, in humic soil, ascending to ca 500 m alt. 6. Gymnosiphon neglectus JoNKER, Monogr. (1938) 7S: Stem simple or branched, 7!/2-11 cm, bearing 1 or few flowers or a capitate inflorescence, consisting of contracted cymes. Leaves lanceolate-ovate, acu- minate, keeled, 1—2!/2 mm. Bracts to 5 mm. Flow- ers subsessile. Perianth dirty white; tube avout 2mm; limb about 2mm. Outer perianth lobes ovate, acuminate; lateral lobes crenate. Inner ones small, linear, acute. Stigmas funnel-shaped, curved, dorsally cristate. Ovary ovoid, about 2 mm. Cap- sule nearly globose, crowned by the 2mm long persistent part of the tube. Distr. Malaysia: Java (Preanger Regencies and Semarang). Ecol. Moist forests, ascending to 1000 m alt. 7. Gymnosiphon pauciflorus SCHLTR. Bot. Jahrb. 49 (1913) 102; Jonker, Monogr. (1938) 176. Stem simple, colourless, 2!/2-9!/2 cm, bearing 1-3 sessile or subsessile flowers. Leaves ovate, acumi- nate, 1-1'/2 mm. Bracts to 3 mm. Perianth whitish to purplish; tube about 4 mm; limb very short. Outer perianth lobes ovate, acute, about I-1!/2mm; lateral lobes entire. Inner ones minute. Connective not apiculate, forked at the top. Style branches rather long, each bearing an ovoid, in transverse section somewhat triangular, stigma. Ovary obo- void, about 2 mm. Capsule ovoid, to 3!/2 mm; per- sistent part of the tube 1!/2-2!/2 mm. Distr. Malaysia: New Guinea (Kani Mts), once collected. 3. THISMIA GRIFFITH, Proc. Linn. Soc. 1 (1844) 221; JonKER, Monogr. (1938) 42, 227. Saprophytic, fleshy herbs. Underground part in the Malaysian species coralliform or vermiform and creeping. Stems usually short, seldom branched. Leaves small, scale-like. Below the flowers | or more bracts, sometimes forming an involucre. Flowers actinomorphic or, sometimes, zygomorphic, urceolate to campanulate. Perianth lobes 6, occasionally free and of equal length and size, or inner ones larger, sometimes inner lobes connivent at the apex or connate in the apical part, then forming an erect mitre with 3 holes, in that case outer lobes very small. Stamens 3, free or, usually, stuck together to an anther tube, hanging at an annulus in the perianth throat. Filaments short, ribbon-shaped. Style thick, short, cylin- drical or conical, persistent, bearing at its apex 3 simple or bilabiate stigmas. Ovary obconical or obovoid; the 3 placentas inserted at the bottom or parietally in the basal part of the ovary. Fruit fleshy, cup-shaped, crowned by the persistent, fleshy, basal ring of the perianth tube and the style with the stigmas. Distr. 24 species, in tropical America (Sect. Myostoma and Ophiomeris), tropical Asia (Sect. Euthis- mia and Sarcosiphon), Chicago area, New Zealand and Tasmania (Sect. Rodwaya). 22 FLORA MALESIANA [ser. I, vol. 4! KEY TO THE SPECIES 1. Inner perianth lobes free, spreading or erect. Underground part vermiform, creeping 2. Perianth lobes equal in length and size 3. Flowers zygomorphic, geniculate 3. Flowers actinomorphic. 4. Stems several, flowers 4-6 ina raceme . . 4. Stem simple; flowers usually 1—3, terminal. 5. Leaves and bracts beset with distinct, prominent, blunt processes . 5. Leaves and bracts without processes. 6. Perianth lobes lanceolate, acute to acuminate, flat , 6. Perianth lobes triangular at the base, tapering into long, filiform tentacles. (Sect. Euthismia SCHLTR.) . (SussectT. Odoardoa SCHLTR.) 1. T. chrysops 2. T. racemosa 3. T. grandiflora 4. T. fumida 7. Perianth tube with horizontal bars inside. Stigmas lanceolate. 8. Anthers provided with 3 thick-filiform appendages at the free apical margin. Perianth yellow- ish in the basal part, bright orange-yellow in the upper part and limb. Tentacles bright orange-red at the base. Perianth lobes with tentacles up to 10 mm 5. T. aseroe 8. Anthers with 1 thick-filiform, median appendage and 2 lateral, short teeth at the free apical margin. Perianth white with 6 ochre-brown streaks; lobes with tentacles c. 20 mm 6. T. alba 7. Perianth tube without bars. Apical margin of the anthers with 2 teeth, each bearing a globose body at the top. Stigmas funnel-shaped with prominent margin . Inner perianth lobes larger . 7. T. ophiuris (SUBSECT. Brunonithismia JONK.) H Perianth very zygomorphic, bilabiate. Upper lip ‘fleshy, bent over the opening of the tube 9. Flowers actinomorphic. 8. T. labiata 10. Inner perianth lobes simple. Tube with prominent horizontal bars inside. 11. Anthers with 3 distinct teeth at the free apical margin, each tooth Be a stiff hair. Outer perianth lobes broadly ovate, obtuse, erect . 11. Anthers slightly dentate apically. Outer perianth lobes short, ear-shaped . . 9. T. javanica . 10. T. arachnites 10. Inner perianth lobes consisting of 3 parts. Basal part erect, short, bearing the transverse part, hamate at the base and broadened at the apex. Third part awl-shaped, inserted on the broad apex of the second part. Perianth tube without bars 11. T. neptunis 1. Inner perianth lobes connected at the apex to an erect mitre with 3 holes. Underground part coral- liform (Sect. Sarcosiphon (Bu.)JONK.) 12. Inner perianth lobes linear, connate at the tips, "forming a mitre ‘with large holes. Anthers ciliate in the basal part, toothed at the apex. 13. Apical margin of the anthers provided with 2 constricted in the middle, below the thecae teeth, each bearing a stiff hair. Anthers slightly 12. T. clandestina 13. Apical margin of the anthers provided with 3 teeth, Each. bearing a Sif hair. Anthers constricted at the base, just above the thecae . 13. T. episcopalis 12. Inner perianth lobes spathulate, connate to a | mitre “with rather small holes. Anthers not ciliate, truncate at the apex . 1. Thismia chrysops RipL. Ann. Bot. 9 (1895) 323; JONKER, Monogr. (1938) 237. Stem usually simple and 1-flowered, about 15 cm. Leaves and bracts linear-lanceolate, acute more or less imbricate, up to 4 mm. Perianth tube geniculate; part below knee c. 3 mm, pink with longitudinal striae; the c.5 mm long, upper part and limb chocolate-brown; perianth mouth yellow. Perianth lobes lanceolate, about 7 mm, tapering to filiform tentacles. Annulus prominent, slightly 6-lobed. Anthers quadrangular, provided with a thick hair on both sides of the free, apical margin and a broad, wing-like appendage, inserted on the midline of the connective. Stigmas oblong, bifid. Fruit stalk elongate. Distr. Malaysia: Malay Peninsula (Malacca, Mt Ophir), once collected. 2. Thismia racemosa RIDL. J. Str. Br. R. As. Soc. 69 (1915) 13; JoNKER, Monogr. (1938) 238. Stems several, occasionally branched. Leaves linear, acuminate. Flowers 4—6in a raceme; pedicels 14. T. crocea 1—-1!/2 cm long. Perianth lobes short, triangular- ovate, blunt. Annulus prominent. Tube cylindrical, about 6 mm long. Distr. Malaysia: Malay Peninsula (Pahang), once collected. 3. Thismia grandiflora Ript. Ann. Bot. 9 (1895) 324; JoNKER, Monogr. (1938) 239. Stem simple, 1-flowered, 3 cm, provided with 2 basal, opposite, scale-like lanceolate leaves, about 5 mm, and 2 apical bracts, of the same shape and size as the basal leaves; both leaves and bracts beset with stiff, terete, blunt processes on the back. Perianth urceolate; tube pink with longitudinal striae, 8 mm. Lobes patent, ovate-triangular in the basal part, tapering at the apex to filiform tenta- cles. Annulus prominent. Anthers not or scarcely stuck together, provided with 2 lateral teeth at the free apical margin and a wing-like appendage in- serted at the middle of the connective. Stigmas lanceolate, bifid, acute, papillose. Ovary about 4mm, obovoid, truncate. Dec. 1948] BURMANNIACEAE (Jonker) 23 Distr. Malaysia: Malay Peninsula (Johore), once collected. 4. Thismia fumida Rint. J. Str. Br. R. As. Soc. 22 (1890) 338; JoNKER, Monogr. (1938) 240. Stem slender, conspicuous, unbranched, about 10 cm, bearing 1—2 flowers. Leaves very small, ap- pressed, lanceolate, acute. Flowers up to 1 cm. Perianth lobes lanceolate, acute, greenish-grey, constricted above the ovary and broadened below the limb, white with pink stripes. Annulus pro- minent. Ovary obconical. Capsule cup-shaped, rib- bed and scabrid, crowned by the crenulate, basal ring of the perianth. Distr. Malaysia: Malay Peninsula (Singapore and Selangor). 5. Thismia aseroe Becc. Malesia 1 (1877) 252; JONKER, Monogr. (1938) 240.—Fig. 3. Stem simple or, sometimes, branched, 1- or 2- flowered, up to 8!/2cm high. Leaves few, lanceolate, obtuse, to 4mm long. At the base of the flowers an involucre, consisting of lanceolate bracts. Peri- anth obconic-campanulate, dirty-yellow in the basal part, bright orange-yellow in the upper part and in the limb. Perianth tube about 11 mm; the basal 5 mm inside with transverse bars. Lobes tri- angular, 3 mm _ long, ending in bright orange tentacles, red at the base, 6mm long. Annulus prominent. Anthers with 3 short thick-filiform ap- pendages at the free apical margin; in the lower part of the anther, inserted at the middle of the connective, a broad, dorsal quadrangular wing, wider than the anther. Thecae oblong, in the basal part of the anther; in the apical part 2 nectaries on the line of junction of one connective with the next. Ovary obovoid, 3 mm. Stigmas narrow-lan- ceolate, rather long, acute. Capsule ribbed, about 5 mm. Fruit stalk lengthening about 5-7 mm above the involucre. Seeds ellipsoid, ribbed. Distr. Malaysia: Malay Peninsula (Singapore and Perak). Ecol. In humic forests. 6. Thismia alba HOLTTUM, ms. A Th. aseroe differt antheris singulis munitis una tantum appendice mediana crassi-filiformi, porro margine apicali libera instructa dentibus 2 late- ralibus brevibus. Perianthium album signatumstriis 6 longitudinalibus ochraceo-brunneis; perianthii lobis tentaculis + 2 cm longis praeditis. Stem simple, 1- to 3-flowered, up to 10cm. Leaves few, 3-4'/2 mm long, lanceolate, acute or acuminate. At the base of the flowers an involucre, consisting of lanceolate, acute bracts. Perianth ob- conic-campanulate, white with 6 thin ochraceous- brown streaks, leading down from each perianth lobe, alternating with 6 thin yellow lines. Perianth tube about 10 mm, the basal part with transverse bars inside. Lobes triangular, 3-4 mm long, pale- yellow at the base, terminated by white, tentacles about 15 mm long. Annulus prominent, bright yellow. Anthers with 1 thick-filiform, median ap- pendage and 2 lateral short teeth at the free apical margin; in the lower part of the anther, inserted at the middle of the connective, a broad, dorsal, quadrangular wing, wider than the anther. The- cae oblong, in the basal part of the anther; in the apical part 2 nectaries on the line of junction of one connective with the next. Ovary semi-globose, about 2 mm. Stigmas lanceolate, retuse, papillose. Capsule obconical, about 6 mm. Distr. Malaysia: Malay Peninsula (Pahang). 7. Thismia ophiuris BEcc. Malesia I (1877) 252; JONKER, Monogr. (1938) 242. Stem 2-6 mm, simple or branched, 1- or 2-flow- ered. Leaves lanceolate, obtuse, to 4mm long. Below the flowers an involucre of several, lance- olate, 3-4 mm long and 1 mm wide bracts. Perianth urceolate, yellowish brown. Tube about 9 mm; lobes triangular, terminated by long, filiform ten- tacles, about 13 mm. Annulus broad and thick. Insertion of the stamens broad, then narrowed into a ribbon-shaped filament and again broadened into the quadrangular anthers. Apical free margin of the anthers provided with 2 teeth, each with a globose body at the top. Ovary ovoid, about 5 mm. Style bearing 3 sessile, funnel-shaped, circumval- lated stigmas. Capsule ribbed; seeds oblong with longitudinal ribs; funicles about the same length as the seeds. Distr. Malaysia: Borneo (Sarawak & Br. N. Borneo). 8. Thismia labiata J.J.S. Bull. Jard. Bot. Btzg III, 9 (1927) 220; JonKER, Monogr. (1938) 44, 243. Stem simple, 22 mm long. Leaves ovate, acute, appressed, 1!/2mm. Flowers with an involucre of 3 ovate-lanceolate, acute, 5'/2 mm long bracts. Pe- rianth urceolate in the basal part, bilabiate-zygo- morphous in the upper part. Outer perianth lobes 2'/2 mm, broad-ovate at the base, rounded, witha subulate appendage inserted below the top. Inner lobes linear to filiform, subulate, 5mm. A thick, fleshy upper lip bent over the perianth mouth; on the back of the upper lip 1 inner and 2 outer peri- anth lobes. The other 2 inner lobes between the 2 lips. The third outer lobe inserted on the middle of the lower lip. Stamens rounded and ciliate at the free, apical margin; thecae elongate; outer side of the stamen provided with scattered hairs. The 3 stigmas connate to a capitate, 3-lobed stigma. Ovary obconical. Distr. Malaysia: Sumatra (Eastcoast Res.) once collected. 9. Thismia javanica J.J.S. Ann. Jard. Bot. Btzg 23 (1910) 32; JoNKER, Monogr. (1938) 245.—Fig. 11. Stem simple or branched, up to 12 cm, 1- to 5- flowered. Leaves ovate or lanceolate-ovate, obtuse, 3 mm. At the base of a flower an involucre of 3 bracts. Perianth tube 7 mm, urceolate, whitish with 12 longitudinal, orange stripes, inside with longi- tudinal bars connected by many transverse bars. Outer perianth lobes obtuse, ovate; inner ones triangular, terminated in up to 3 cm long, filiform tentacles. Anthers 3-toothed at the free apical mar- gin; each tooth terminating in a hair. On the outer side of the anther, inserted in the middle, a qua- 24 FLORA MALESIANA [ser. I, vol. 4! drangular appendage, wider than the stamen. Margin of the appendage strigose. Style orange- coloured; stigmas sessile, ovate, truncate. Ovary obovoid, 3 mm. Capsule orange-coloured, about 6 mm. Seeds ellipsoid. Fig. 11. Thismia javanica J. J. S. Doengoes Iwoel, < 3/2. (LIEFTINCK) Distr. Malaysia: Sumatra and W. Java. Ecol. Shade of forests, on humus, 1000 m alt. Vern. Angkrek rambut (Java). Notes. Perhaps conspecific with the following species. below 10. Thismia arachnites RipL. Journ. Str. Br. Roy. As. Soc. 44(1905) 197; JoNKER, Monogr. (1938) 247. Stem simple, 1—71/2 cm, bearing 1-3 flowers. Leaves few, lanceolate, acute, about 5 mm. At the base of the flowers an involucre, consisting of ovate-lance- olate, acute, 5~7 mm long bracts. Perianth urceolate to obconical, about 8 mm long, inside with longi- tudinal bars, connected by many transverse bars; tube transparent, white with 6, vertical, red streaks in the apical part. Perianth lobes pale red. Outer lobes very short, about 4 mm, ear-shaped; inner ones triangular, about 1 mm, terminating in up to 3 cm long, thin, filiform tentacles. Annulus prominent, yellow. Anthers slightly 3-toothed at the apical free margin, the lateral teeth somewhat larger than the median one, each tooth terminating in an indistinct, very thin, fragile hair. On the outer side of the anther, inserted in the middle, a quad- rangular strigose appendage, wider than thestamen. Style thick, conical; stigmas lanceolate, obtuse. Fruit stalk lengthening above the involucre. Distr. Malaysia: Malay Peninsula (Perak & Pahang). Note. Closely related to the preceding species, perhaps conspecific. 11. Thismia neptunis BEcc. Malesia I (1877) 251; JONKER, Monogr. (1938) 43, 243. Stem 4-25 mm, simple, 1-flowered, beset with few, lanceolate, acute, about 3 mm long /eaves. At the base of the flower an involucre of 3 lanceolate, acute, 46mm long bracts. Perianth tube urceolate, with 6 longitudinal stripes, 6 mm. Outer perianth lobes simple, recurved, filiform with triangular base, 4!/2 mm. Inner ones erect, about 15mm, con- sisting of an erect, short, basal part; a transverse part with hamate base and a broadened, rounded apex and, inserted on the apex of the transverse part, anerect, long, awl-shaped part. Annulus prom- inent. Anther quadrangular, 3-toothed at the free apical margin. Stigmas lanceolate, acute. Ovary obovoid, truncate, 1!/2 mm. Distr. Malaysia: Borneo (Sarawak, Mt Mat- tang), once collected. 12. Thismia clandestina (BL.) Mia. Fl. Ind. Bat. 3 (1855) 616; JoNKER, Monogr. (1938) 252.—Sarco- siphon clandestinus BL. Mus. Bot. Lugd. Bat. 1 (1849) 65. Stem up to 5cm, 1—2-flowered. Leaves appressed, acuminate, to 5 mm. At the base of the flower 3 bracts. Perianth tube urceolate, about 2!/2 mm, greenish-grey, with 12 longitudinal, brownish- black stripes. Outer perianth lobes almost absent, inner ones connate to a 2!/2 mm long, acuminate mitre. Annulus prominent, 6-lobed. Margin of the filaments and upper part of the anthers with short hairs. Thecae oblong, inserted on the margins of the anthers. Free apical margin of the anthers with 2 teeth, tapering to stiff hairs. Inserted on the midline a large, wing-like appendage, provided with bundles of hairs on the angles. Stigmas ovate, bilobate, papillose, whitish. Ovary obovoid, about 3 mm. Funicle about the same length as the ovules. Capsule papillose, about 5 mm. Distr. Malaysia: W. Java. Ecol. In humus of forests, ascending to ca 1000 m alt. 13. Thismia episcopalis (BECc.) F. v. MUELL. Pap. & Proc. R. Soc. Tasm. for 1890 (1891) 235; JONKER, Monogr. (1938) 46, 253.—Geomitra episcopalis Becc. Malesia 1 (1877) 250.—Bagnisia episcopalis ENGL. Pfl. Fam. 2, 6 (1889) 48.—Sarcosiphon episco- palis Scutr. Notizbl. 8 (1921) 38.—Fig. 4. Stem simple or branched, 1-8 flowered, up to 19cm. Leaves appressed, ovate, acute, 2-5 mm. Perianth tube urceolate, yellow with black stripes, 6-9 mm. Outer perianth lobes almost lacking; inner connate to a slightly acuminate, about 5 mm long mitre. Filaments constricted. Thecae diver- gent. Margin of the anthers and the winglike ap- pendage hairy, apical part of the anther, below the insertion of the appendage, darker coloured; free apical margin 3-toothed, each tooth terminated by a stiff hair. Stigmas bilobate. Ovary obovoid, 3 mm. Funicles about as long as the ovules. Capsule ribbed, about 3 mm. Fruit stalk lengthened. Dec. 1948] BURMANNIACEAE (Jonker) 25 Distr. Malaysia: Borneo (Sarawak, Br. N. Borneo). Ecol. In humus of forests, ascending to ca 1700 m alt. 14. Thismia crocea (BeEcc.) J.J.S. Nova Guinea 8,1(1909) 193; JonKER, Monogr. (1938) 44, 251. —Bagnisia crocea BEcc. Malesia 1 (1877) 249.— Thismia versteegii J.J.S. Nova Guinea 8,1 (1909) 193.—Sarcosiphon croceus SCHLTR. Notizbl. 8 (1921) 38.— Sarcosiphon versteegii SCHLTR. Notizbl. 8 (1921) 38. Stem simple, 1—3-flowered, about 6 cm. Leaves appressed in the basal part, lanceolate, acute, to 6 mm. At the base of the flowers 3 ovate lanceolate, acute bracts. Perianth tube urceolate, ribbed, red- dish-brown in the upper part, yellowish-orange in the middle and white at the base, about 6 mm. Outer perianth lobes broad, short, rounded; inner ones connate to a thick, 2mm long mitre with 3 narrow holes and 3 prominent midribs. Annulus slightly 12-lobed. Anthers quadrangular, not hairy; thecae oblong, parallel; inserted in the apical part of the anther a broad appendage with curled mar- gins. Stigmas ovate, obtuse, papillose. Ovary light reddish-brown, about 2 mm. Funicles as long as the ovules. Fruit ribbed, obovoid. Fruit stalk thickened and lengthened after flowering to 16 mm above the bracts. Distr. Malaysia: West New Guinea. Notes. In Perak (Malay Peninsula), RIDLEY observed a Thismia, described by him in Mat. Fl. Mal. Pen. 2 (1907) 75, as Bagnisia crocea var. brunnea. This specimen was apparently not preserved; it is highly improbable that it belongs to T. crocea. 4. GEOMITRA BeccarI, Malesia 1 (1877) 250; JONKER, Monogr. (1938) 46, 254. Underground part unknown. Stem beset with scale-like /eaves. Flowers rather large, with an involucre at the base. Tubular part of the perianth urceolate. Outer perianth lobes free, very small. Inner ones connate at the top to an erect mitre with 3 holes, crowned by 3 apical, long, thick-filiform, erect, clavately swollen appendages. Basal ring of the perianth tube thickened, persistent on the fruit. Throat margin of the perianth thickened to a 6-lobed annulus. Stamens 6, hanging at the annulus; anthers stuck together to a tube. Style short, cylindrical, fleshy, bearing 3 erect stigmas. Ovary with 3 stalked placentas; funicles short. Capsule cup-shaped, crowned by the persistent, basal perianth ring and the style. Distr. One species, known only from Borneo (Sarawak). 1. Geomitra clavigera BEcc. Malesia 1 (1877) 251; JONKER, Monogr. (1938) 46, 255.—Thismia clavi- gera F. v. MUELL. Vict. Nat. (1890) 235.—Sarco- siphon clavigerus SCHLTR. Notizbl. 8 (1921) 39. Stem simple, up to 12 cm, bearing about 3 flow- ers. Leaves lanceolate, acuminate or acute, 2-6 mm. Bracts lanceolate, acuminate, 6-7 mm. Perianth 3—5 mm, hooked at the apex. Filiform appendages 8-12 mm long. Anthers quadrangular; free apical margin with 3 teeth, each bearing a stiff, trans- parent hair. Anther tube about 4mm. Stigmas lanceolate, bilobate; lobes acute. Ovary obovoid, truncate, about 3 mm. Distr. Malaysia: Borneo (Sarawak), once col- tube about 9mm. Outer perianth lobes erect, lected. broadly triangular, about 1mm. Mitre about 5. SCAPHIOPHORA SCHLTR. Notizbl. 8 (1921) 39; JoNKER, Monogr. (1938) 46, 256. Roots coralliform. Stem provided with scale-like Jeaves; at the base of the flower an involucre. Perianth tube urceolate. Outer perianth lobes small; inner ones narrow in the basal part, broadened at the apical part, connate to an erect mitre with 3 holes in the basal part. Mitre crowned by a long, stiff column, bearing at the top 3 lobes. Stamens 6, hanging, inserted at an annulus in the perianth throat. Filaments ribbon-shaped. Anthers stuck together to an anther tube; each anther provided with a wing-like appendage, inserted in the middle and broader than the anther. Placentas stalked; stalks inserted peripherically at the bottom of the ovary. Basal perianth ring and style persistent on the fruit. Distr. Two species, one in New Guinea, the other in the Philippines. 26 FLORA MALESIANA [ser. I, vol. 4', Dec. 1948] KEY TO THE SPECIES 1. Flowers 3-6!/2cm long (without column). Column 1!/2-6 cm long, at the apex broadened to 3 fleshy, connate lobes . 1. S. gigantea 1. Flowers 1 cm long (without column). Column 5 mm long, bearing at the apex 3 cup-shaped bodies . 1. Scaphiophora gigantea JONKER, Monogr. (1938) 257.—Fig. 2. Stem 4-10!/2 cm, partly subterranean. Leaves lanceolate, acute, 2-4mm, the lower ones keeled. Bracts ovate, lanceolate, acute, about 18 mm. Peri- anth tube 15-21 mm, pale rose-coloured with yellow veins, reticulate below the inner perianth lobes. Outer perianth lobes ear-shaped. Mitre 5-9 mm long, orange to yolk-yellow. Stamens about 7mm. Anthers prominently nerved; free apical margin provided with 3 median and 2 lateral teeth; each bearing a stiff, transparent hair. Appendix of the anther greenish-blue; lateral margins bearing 3 bundles of short hairs; apical margin pilose. Thecae divergent, ovate. Style truncate-conical. Stigmas sessile, obovate, 2-lobed, papillose outside and in the upper part inside. Fruit cup-shaped. Placentas connate at the apex, stalked; stalks about the same length as the placentas. Distr. Malaysia: Philippines (Luzon), twice collected. 2. S. appendiculata 2. Scaphiophora appendiculata (SCHLTR.) SCHLTR. Notizbl. 8 (1921) 39; JoNKER, Monogr. (1938) 259. —Thismia appendiculata SCHLTR. Bot. Jahrb. 55 (1918) 202. Stem 15-20 mm high, partly subterranean, usu- ally 1-flowered. Leaves ovate to lanceolate, 2—3 mm. Bracts lanceolate, acute, about 5mm. Perianth tube 6mm, yellowish white in the lower part. Outer perianth lobes small, ear-shaped. Mitre 3-6 mm, orange-coloured. At the base of each peri- anth lobe, on theinner side, a glandular, bowl-shaped body. Column + broadened towards the apex, bearing 3 thick, fleshy, cup-shaped bodies. Stamens about 3 mm. Appendage of the anther crenulate at the apical margin. Thecae divergent. Style trun- cate-conical, 11/2 mm. Stigmas sessile, obovate, 2- lobed, 1 mm. Ovary 3!/2 mm. Placentas stalked, above the fertile part suddenly narrowed again into a filiform, apical appendage. Placentas attach- ed to the bottom of the ovary by the stalks and to the roof by the apical appendages. Distr. Malaysia: Northeast New Guinea, once collected. SPHENOCLEACEAE (H. K. Airy Shaw, Kew) Mart. ex LINDL. Nat. Syst. ed. 2 (1836) 238; DC. Prod. 8 (1939) 548; Wicut, III. Ind. bot. 2 (1850) 115; Mia. FI. Ind. Bat. 2 (1857) 569; Borss. Fl. Or. 3 (1875) 963. Annual (?)laticiferous herbs, with the habit of Phytolacca. Stem erect, somewhat succulent. Leaves spirally arranged, simple, entire, exstipulate. Inflorescences termi- nal, densely spicate, acropetal. Flowers subtended by a bract and two bracteoles, _ bisexual, actinomorphic. Calyx tube adnate to the ovary; segments 5, united below, imbricate, connivent, persistent. Corolla campanulate-urceolate, perigynous; lobes 5, imbricate. Stamens 5, epipetalous, alternating with the corolla lobes; filaments short; anthers rounded, 2-locular, dehiscing longitudinally. Ovary semi-inferior, _2-locular; style short, stigma capitate; ovules ~, attached to large spongy stipitate _axile placentas. Capsule cuneate-obconic, 2-locular, membranous, circumscissile; seeds ~, minute, oblong, rugose-costate, albumen very scanty or none (?); embryo axile, straight, subterete. Distr. Mono-generic, almost pantropical. Ecol., Uses, Vern., see below under S. zeylanica. Notes. The maintenance of Sphenocleaceae as a separate family is abundantly justified; there is no evidence of affinity with Campanulaceae, with which it has hitherto been associated. The habit resembles that of Phytolacca, and the anatomy shows several significant features occurring in members of the Phytolaccaceae and related families. Other characters suggest Primulaceae, and provisionally it is sug- gested that the family represents a ‘half-way house’ between the families mentioned. From the Centro- spermae it deviates in the semi-inferior ovary, gamopetalous corolla and straight embryo, and from the Primulaceae principally in the alternipetalous stamens. A separate note on the classification will be published in the Kew Bulletin. 1. SPHENOCLEA GAERTN. Fruct. 1 (1788) 113, t. 24, f. 5; Mig. FI. Ind. Bat. 2 (1857) 569; B. & H. Gen. Pl. 2 (1876) 560; BAILL. Hist. Pl. 8 (1886) 327, 362, f. 158-161; SCHONLAND, in E. & P. 4, 5 (1889) 60; BoeRL. Hand. 2, 1 (1891) 257. For characters see family description. Distr. Two species, one pantropical, one endemic in W. Africa. 1. S. zeylanica GAERTN. Fruct. /.c.; Bl. Bijdr. 16 nivent. Corolla whitish, 2!/2-4 mm long, caducous, (1826) 1138; Moritz, Syst. Verz. (1845-6) 66; segments ovate-triangular, obtuse or acute, united BLANCO, FI. Filip. ed. 2(1845) 62, ed. 3, 1(1877) 117, t. 143; Mia. /.c.; F.-VILL. Nov. App. (1880) 121; K. & G. Mat. Fl. Mal. Pen. no 16 (1905) 57; Koorp. Exk. Fl. 3 (1912) 301; Merr. Fl. Man. (1912) 462; Sp. Blanc. (1918) 374; En. Philip. 3(1923) 588; Rpt. Fl. Mal. Pen. 2 (1923) 204; BAcker, Onkruidfl. Jay. Suik. (1931) 742; OcHsE & BAKH. V.D. BR. Veg. D. E. I. (1931) 93, f. 55, 349.—Pongatium spongi- osum BLANCO, FI. Filip. (1837) 86.—Reichelia palustris BLANCO, /.c. 220; ed. 2 (1845) 155; ed. 3, 1 (1877) 277, t. 143.—Fig. 1. Roots long, cord-like. Stem hollow, 7-150 cm. Leaves oblong to lanceolate-oblong, attenuate at both ends, acute or obtuse, glabrous, 2!/2-12!/2 by '/2-5 cm; petiole 3-30 mm. Spikes 3/4~7!/2 cm long, cylindric; peduncle 1-8 cm. Bracts and bracteoles + spatulate, the green apices arched over the calyx before and after anthesis. Flowers crowded, rhomboid or hexagonal by compression, sessile, wedge-shaped below, attached longitudinally to the rachis by a linear base. Calyx segments deltoid- semicircular, obtuse, ultimately accrescent and con- slightly more than half-way, connivent. Stamens inserted half-way up tube of corolla, filaments slightly dilated at base. Ovary obovoid, 2!/2 mm long, apex broad, free, truncate. Capsule 4-5 mm in diam., dehiscing below the calyx segments which fall with the lid, leaving the scarious persistent base. Seeds yellowish-brown, + '/2 mm long. Distr. Trop. America (introduced), trop. Africa (incl. Madagascar) (indigenous; cf. BENTH. in Journ. Linn. Soc. Bot. 15 (1875) 13), SW. Persia to Turkestan, India and Formosa (prob. intro- duced), in Malaysia (prob. introduced): Malay Peninsula (scarce, mainly in the prov. Kedah and Wellesley), 72Sumatra, Philippines (Luzon, Biliran, Negros), Java, Bali, SW. Celebes and Timor. Ecol. A weedy annual occurring in almost any kind of damp ground at low alt. up to 350 m: river banks and dry riverbeds, damp marshy or periodic- ally inundated depressions, seasonal swamps, sides of ponds, ditches, and stagnant water generally, especially rice-fields, both in continuously rainy and in seasonal climates. Almost every flower on 28 FLORA MALESIANA [ser. I, vol. 4', Dec. 1948] every inflor. sets fruit; only one or two flowers are open at once on any one head. In Malaysia never gregarious, nor growing on mud of tidal creeks, as in Africa. Uses. In Java young plants and tips of older plants are steamed and eaten with rice; they have a slightly bitter taste; leaves are sold under the name goenda padi. Vern. Java: goenda, M, J, Sd, g. rawah, g. lalab, g. padi, g. sapi, Sd, goendha, Md, gondo, J; Bali: gonda; Celebes: gangang karaéng, Mk., gonra, Mk, Bg; Philippines: mais-mais (Panay, Bisaya), silisi- lihan (Tagalog); the Javanese names are also ap- plied to the superficially similar Hydrophyllaceous Hydrolea zeylanica (L.) VAHL. Notes. The plant is described as laticiferous but METCALFE reports that ‘typical laticiferous ca- nals are absent from the phloem, although occa- sional elongated cells have been observed in this tissue with granular contents which may represent coagulated latex’. Miss M. C. VREEDE, Anatomist in the Treub Lab., Buitenzorg, Java, reported, July 6, 1948, that in fresh material she could find neither milky juice nor laticiferous elements. Fig. 1. Sphenoclea zeylanica GAERTN. X 1/4. A rich-flowering individual. NYSSACEAE! (J. Wasscher, Groningen) 1. NYSSA LINNE, Sp. Pl. (1753) 1058; Gen. Pl. ed. 5 (1754) 478; WasscHer, Blumea 1 (1935) 343.—Agathisanthes & Ceratostachys Bl. Bijdr. (1825) 644; Mig. FI. Ind. Bat. 1, 1 (1856) 838.—Agathidanthes HASSsK. Cat. Hort. Bog. (1844) 254.—Daphniphyllopsis Kurz, J. As. Soc. Beng. 44, II (1875) 201. Dioecious trees or shrubs. Leaves simple, scattered. Stipules 0. Flowers uni- sexual, often in heads, in the axils of a bract and with 2 bracteoles. d: in axillary heads or short racemes; calyx entire or 5-toothed; petals 5, imbricate, often small, alternate with the calyx; stamens 8-16 in 2 alternating whorls; anthers small, dorsifixed with lateral lengthwise slits; disk pulvinate; style rudimentary. 9: soli- tary, axillary or in 2-10-flowered heads; ovary inferior, l1-locular, connate with the 5-toothed or entire calyx; petals 5-8 often minute; stamens of inner whorl partly sterile, both petals and anthers soon dropping; style with 2 appressed later diver- gent often torulose branches stigmatose on their inside, brittle, often deficient in the herbarium. Ovule 1, hanging from the apex of the cell, anatropous with 2 integu- ments. Fruit drupaceous ovoid to oblong. Distr. Ca 6 spp., 4 in Atlantic N. America, 1 in China, 1 from India to W. Malaysia. Ecol. The American spp. mostly in swamp forests, the Asiatic one not so. Notes. The flowers are often deficient in the herbaria. The polymorphy of N. javanica suggests that perhaps more than one species is present in Malaysia. 1. Nyssa javanica (BL.) WANG. Pfl. R. 41 (1909) 15; WASSCHER, Blumea 1 (1935) 344.—Ceratostachys arborea BL. Bijdr. (1825) 644; Mra. FI. Ind. Bat. 1, 1 (1856) 839.—Agathisanthes javanica BL. Bijdr. (1825) 645; Mia. FI. Ind. Bat. 1, 1 (1856) 839.— Agathidanthes javanica Hassk. Cat. Hort. Bog. (1844) 254.—Nyssa sessiliflora Hook. f. & Th. Gen. Pl. 1 (1867) 952.—Ilex daphniphylloides Kurz, J. As. Soc. Beng. 39, II (1870) 72.—Daphniphyllopsis | capitata Kurz, l.c. 44, II (1875) 201; For. Fl. Burm. 1 (1877) 240.—Nyssa arborea Koorp. Exk. FI. Jav. 2 (1912) 731.—Nyssa bifida CraiB, Kew Bull. (1913) 69.—Fig. 1. Dioecious tree up to 40m, 30-100 cm diam., clear bole 13-23 m, buttresses mostly absent. Twigs tomentose, glabrescent. Leaves rather den- sely set, oblong-lanceolate to obovate, rarely sub- ovate, base acute, apex abruptly acuminate, cori- aceous, entire, sparsely hairy to tomentose on midrib and nerves beneath, further glabrous, 5—23 by 2!/2-8 cm; in seedlings the Ist pair of leaves is opposite; nerves 8-11 pairs; petiole 1-3!/2 cm long, flat or slightly sulcate, hairy or glabrous. Flowers pallid, in pedunculate nearly globose axillary heads 12-18mm diameter; peduncles flat- tened towards the apex 3/4—5 cm long, their apex 2-5 mm broad, glabrous or hairy, ca halfway with 1—2 sessile small acute bracts 3-4 by 1 mm. Re- ceptacle globose to ellipsoid, flattened, 2-3 and 4-5 mm. Flowers enveloped by 1 bract and 2 half- way connate bracteoles, all broad-ovate, sericeous- ciliate, 2—2'/2 by 11/2-3 mm, in Q persistent.— Fig. 1. Sarcosperma uittienii H. J. L. a. flowering branch nal and cross-section. the corolla and calyx within 34 FLORA MALESIANA [ser. I, vol. 41, Dec. 1948] Vern. Not constant, few noted. Notes. It is probable that more specimens are hidden among indeterminates in several families. 2. Sarcosperma uittienii H. J. LAM, Bull. J. B. B. III, 8 (1926) 19, f. 1, &c.—S. sumatranum UITT. ex Lam, /.c.—Fig. 1. Tree. Leaves fairly opposite, oblong-elliptic to ovate or obovate, both base and apex acuminate, glabrous above glabrescent below; petiole 12-20 mm. Inflor. densely minutely tomentose, generally broadly and laxly paniculate, sometimes almost unbranched, 33/4-131/4 cm long, branches 11/2-63/4 cm long; bracts tomentose, deltoid 1-2 mm long. Flowers fascicled or solitary only known in bud; pedicals 2-4 mm. Calyx densely tomentose, 2!/2 by 2mm. Corolla tube !/2 mm long, lobes obovate, 2-21/2 mm. Staminodes deltoid 1/2 by '/3 mm. Sta- mens ovoid, 1 mm through. Ovary glabrous, 2!/2 by 1!/2 mm. Style 1 mm. Fr. unknown. Distr. Malaysia: only known from Sumatra (Eastcoast Res.). Ecol. Forests, ca 500 m. Fl. June—July. Vern. Only once noted. Notes. Inadequately known. Closely related to S. kachinense (KING & PRAIN) ExELL from Burma & China, and to S. arboreum Hook. f. from India | to China. | Excluded Sarcospermum petasites REINW. eX DE VRIESE, Rein- wardt’s reize (1858) 576=Gunnera macrophylla BL. (Halorrh.). STACKHOUSIACEAE (F. I. Brouwer, Groningen) STACKHOUSIA J. SMITH, Trans. Linn. Soc. Lond. 4 (1798) 218; PAmp. Bull. Herb. Boiss. II, 5 (1905) 912; Brouwer, Blumea 3 (1938) 173; MATTF., in E. & P. ed. 2, 20b (1942) 240. Annual, or perennial herbs with a rhizome. Leaves scattered, entire. Stipules 0 or very small. Racemes terminal. Flowers bisexual, regular, 5-merous, in groups in the axils of bracts. Sepals usually more or less connate, rarely free. Corolla perigynous or almost hypogynous, petals long-clawed, rarely entirely free, usually free at the base, connate in the upper portion of the claws, lobes imbricate spread- ing. Stamens 5, inserted on the margin of the calyx tube, free, usually unequal (2 shortest), included in the corolla tube. Ovary (2-)3(-5) celled, lobed, each cell with | erect ovule. Style with (2-)3(-5) stigmatic lobes, partly sunk in the ovary. Fruit with (2-)3(-5S) one-seeded cocci and a columella. Distr. Ca 19 spp. in Australia, 4 in Tasmania, 1 in New Zealand and 1 in Malaysia, Australia and Micronesia (Palau, Jap). Notes. The family consists next to the genus Stackhousia, the single one by which it is represented in Malaysia, of 2 other monotypic genera, and is practically confined to Australia. It is not directly allied to any other family and has been compared with e.g. Euphorbiaceae, Celastraceae, Sapindales, &c. 1. Stackhousia intermedia F. M. BaILey, Q. Agric. J. 3, 4 (1898) 174; Q. FI. (1899) 264; Pamp. J.c. (1149, cum f. philippinensis; BROUWER, Blumea 3 (1938) 174; STEEN. J. Arn. Arb. 28 (1947) 422. —S. muricata (non LINDL.) auct. plur. quoad Philip. _—St. viminea (non J. SM.) VOLKENS, Bot.Jahrb. 31 (1902) 467; id. var. micrantha LAutB. Nachtr. FI. Deut. Sch. Geb. Siids. (1905) 305.—St. tenuissima var. ramosa STEEN. Nova Guinea 14 (1927) 307.— Fig. 1—2. Erect, glabrous annual, 6-50 cm long. Root fusi- form, up to 5 cm long, 1!/2 mm diam. at the base, attenuate, with fibrous ramifications. Stem gradu- ally attenuate to the almost filiform angular apex, little branched and leafy below, terete, striate, inter- nodes '/2-3 cm long. Leaves linear, sessile, base attenuate, 7!/2-20 by !/s—2 mm, lower obtuse, upper acute to mucronate, nervation absent or midrib visible. Racemes 1—20cm long. Flowers minute yellow, upper groups 1-3 fls and 2 bracteoles, low- er groups with more bracteoles and up to 5 fis. Bracts roundish ovate, strongly acuminate, fim- briate, dentate, 3/s-1 by '/2 mm, membranous ex- cept the midrib. Bracteoles like the bracts but more dentate and less acuminate. Pedicels 3/4—1!/4 mm. Calyx-tube 1!/2mm high, lobes ovate-acuminate, 1/2 mm long, irregularly fimbriate-dentate, margin membranous. Corolla inserted on the margin of the calyx-tube, sympetalous, hypocraterimorphous, tube cylindrical, 2 by '/2 mm, divided into 5 petals in the lower portion over !/4 mm, lobes ovate-ob- long, strongly acuminate, ca 3/4mm long. Fila- ments filiform, 2 shorter ones reaching the middle, 3 longer ones the margin of the corolla-tube; anthers oblong, very obtuse and emarginate at base and apex, 0.6 by 0.3 mm, introrse, dithecic, 4-lo- cular. Ovary subglobose, 0.3—0.4 mm diam. 3-lo- bate, 3-celled. Style straight, 0.4 mm long, with 3 linear stigmas. Cocci 3, roundish ovate, 11/2 by 1 mn, reticulate. Distr. Australia, Micronesia, and Malaysia: Sumatra (Toba-Batak Lands), N. Celebes, Philip- pines (Luzon, Culion, Guimaras), Moluccas (Boe- roe, Ambon, Saparoea), New Guinea, 10—100—300-— 600-1500 m alt.—Fig. 1. Fig. 1. Localities of Stackhousia intermedia BAILEY in Malaysia. Ecol. Lank herb mostly in grassfields, savan- nahs and abandoned fields, in both wet and dry spots, in Sumatra at 600-1400, but in E. Malaysia & Micronesia below 300m, in the Philippines ascending to 1500m. F/. mostly in April-May together with the grasses. Notes. St. tenuissima, virgata, aphylla and mi- crantha Pamp. l.c. are most probably all identical with this species. 36 FLORA MALESIANA _ (ser. J, vol. 4!, Dec. 1948] Fig. 2. Stackhousia intermedia BAILEY, X 1/2, fruits and flowers enlarged. ACTINIDIACEAE (C. G. G. J. van Steenis, Buitenzorg) 1. ACTINIDIA LINDL. Nat. Syst. ed. 2 (1836) 439; B. & H. Gen. Pl. 1 (1862) 177; BENTH. FI. Hongk. (1861) 26; KiNG, Ann. R. Bot. Gard. Calc. 5, 2 (1896) 145, t. 176; E. & paca, 21 (1925) 36. Trailing shrubs or lianas without special organs for climbing, branches rarely flexuose; stem with wide vessels, raphides in the flowering parts: bark often with short linear lengthwise lenticels. Growth in flushes from terminal and axillary buds. Indumentum of stellate or simple hairs. Stipules minute, obsolete, or absent. Leaves simple, scattered, petiolate, serrate or callous-dentate, penninervous, midrib sulcate, veins in cross-bars, veinlets reticulate. Inflor. lateral, often on a common peduncle forked at the apex, cymose, often pseudo-umbellate; bracts 2, at the apex of the peduncle. Flowers mostly white, dioecious (or polygamous), 5(—4)— merous. Sepals distinctly imbricate (rarely valvate), free or subconnate at the base, persistent. Stamens (10—), in 9 fls with short filaments and small sterile anthers; filaments thin, anthers versatile, base divaricate, attached in the middle, reflexed in bud, dehiscing lengthwise. Disc absent. Ovary free, superior, tomentose (or glabrous), (S—) ~-celled; ovules attached on the central axis. Styles free, (5—)s, persistent, elongating after flowering in 9, --clavate, spreading, in d ovary small, with minute styles. Berry glabrous (or hairy), often spotted by lenticels, oblong. Seeds ~, small, biconvex, oblong, immersed in pulp; testa cartilagineous, reticu- late-pitted, dark when dry; albumen copious; integuments |; embryo cylindrical straight, cotyledons short. Distr. Ca 30 spp. from W. Malaysia & Himalaya to Sachalin, Japan and Formosa, centering in China and Japan. Ecol. Forests and forest borders, in the montane zone mostly. Notes. Both Malaysian species appear to be strictly dioecious; the number of ¢ and 9 sheets in A. callosa is about equally large; on Mt Kinabalu only 9 have been found of A. latifolia. The total number of specimens examined is inconsiderable; the species are either rare or little collected being inconspicuous. The genus Actinidia is often included in Theaceae, Dilleniaceae, or even Ericaceae, and it is closely related to Saurauiaceae from which it differs in its trailing or climbing habit, absence of scale-like emer- gences (except in A. strigosa), mostly dioecious fls, ebracteate pedicels, lengthwise dehiscing anthers, numerous styles, and a multilocular ovary. I wish to express my sincere thanks to Mr H. K. Airy SHAW and to Mr M. R. HENDERSON for verifying the MS. of this contribution with the materials preserved at London and Singapore respectively. KEY TO THE SPECIES 1. Leaves either glabrous or subglabrous, or provided with simple pluri-celled hairs. Petals glabrous. Inflorescences short : ees ce Ne Ae, Bah as 1. A. callosa 2. Leaves glabrous or subglabrous ; var. callosa 2. Leaves rather distinctly subtomentose beneath var. pubescens 1. Leaves stellate-tomentose beneath. Petals pubescent on the back. Inflorescences often well-developed. Peduncle 11/2-8 cm 1. Actinidia callosa LINDL. Nat. Syst. ed. 2 (1835) 439, s.l.; K. & V. Bijdr. 3 (1896) 280; BACKER, Schoolfil. (1911) 102; Dunn, J. Linn. Soc. 39 (1911) 405; Koorp. Exk. Fl. 2 (1912) 602; Fl. Tjib. 2 (1923) 179; BAker, J. Bot. (1924) Suppl. 9; STEEN. Bull. J.B.B. IIT, 13 (1934) 174.—See further under var. pubescens. Rambling or trailing shrub or liana up to 30 m, twig-lenticels distinct, wood and inner bark orange. Petiole red s.v., 1-4 cm, blade rather variable in shape ovate-elliptic or obovate, acuminate, midrib 2. A. latifolia red s.v., S—10!/2 by 2!/2-6 cm,sidenerves ca 5—6 pairs rather steeply ascending and substraight, insertion decurrent, margin distinctly serrate or dentate, teeth erect at the end of a vein, apex acuminate, base rounded to subcuneate. Indumentum meagre or absent, consisting of short often somewhat crisped pluri-celled simple hairs. Peduncle, pedicels and calyx thin-tomentose. Peduncle '/4-1!/2 cm, pedicels !/2-1!/4 cm, all thin. Dioecious, flowers white, anthers yellow. Sepals ovate-orbicular, ca 6 by 5 mm. Petals oblique-broad-spathulate, sub- 38 FLORA MALESIANA [ser. I, vol. 4! Fig. 1. Actinidia latifolia (GARDN. & CHAMP.) MERR., habit x ‘/2 (after KING). Dec. 1948] ACTINIDIACEAE (Vv. Steenis) 39 fleshy, margin + irregular, ca 10 by 7 mm. Stamens © in ca 2 rows, filaments subequal, ca 6 mm (in 9 very short); anthers 12/3 by 1 mm, apex subapiculate (in Q sterile, hardly dehiscing); 9 fls unknown to me. Ovary stout cylindric, styles ca 2mm (in d very small, reduced). Berry grey-green, spotted grey or brown, entirely syncarp, oboval to broad-ellip- tic, often oblique, apex often concave, 17—27 by 14-18 mm, base rounded, sepals recurved. Seeds elliptic, 3 by 11/2 mm. Distr. SE. Asia, China, Formosa, in Malaysia: Sumatra, Java. Ecol. Mountain forests, forest borders, 1000- 2040 m, rather rare. Notes. Young shoots edible. Leaves sometimes with raspberry-coloured zoocecidia consisting of crowded-hairy portions. In Java a juvenile shoot was collected with subcordate subglabrous leaves resembling in shape those of A. latifolia. A rather variable species; some of the forms dis- tinguished by DUNN are now taken up as species, wrongly it seems. A. indochinensis MERR. apparently belongs here. var. pubescens DuNN, /.c. 406.—Saurauia tomen- tosa KORTH. nomen ex K. &. V. Bijdr. 3 (1896) 280.—Actinidia pubescens RIDL. J. Fed. Mal. Stat. Mus. 8, 4 (1917) 18.—Leaves 6!/2-11'/2 by 4!/2-6!/2 cm, thinly tomentose beneath. Distr. Assam, in Malaysia: Malay Peninsula (HENDERSON 23436), Sumatra (KORTHALS, FORBES). Notes. Apparently rare, may be confused with A. latifolia. The Sumatra specimen has glabrous twigs, the others hairy ones. The indumentum seems partly caducous. I assume KoORTHALS’s specimens came from Sumatra, not from Java. 2. Actinidia latifolia (GARDN. & CHAMP.) MERR. J. Str. Br. R. As. Soc. 86 (1922) 330.— Heptaca latifolia GARDN. & CHAMP. in HooK. J. Bot. & Kew Gard. Misc. 1 (1849) 243.—Kadsura pubescens MiqQ. Fl. Ind. Bat. Suppl. (1860) 620; Kurz, J. As. Soc. Beng. 45, II (1876) 119, non A. pubescens RIiDL. 1917.—A.championi BENTH. Fl. Hongk. (1861) 26; FINET & GAGN. FI. Gén. I. C. 1 (1907) 28; RIDL. Fl. Mal. Pen. 1 (1922) 206.—A. miquelii KiNG, J. As. Soc. Beng. 59, II (1890) 196, nomen illeg.; Ann. R. Bot. Gard. Calc. 5 (1896) 145, t. 176.— Fig. 1. Rambling shrub or liana to 20m long, twigs dark-coloured s.s., innovations, inflor. and under- surface of the /eaves thinly cinnamon- (s.v. rusty- red-)stellate-tomentose. Petiole 2-4 cm; blade broad-ovate, obovate to suborbicular, 5!/2-11 by 3-9 cm, base reniform-cordate to rounded or cun- eate, apex acuminate, margin subentire with small callous teeth, veins rusty in distinct cross-bars, reticulations below hidden by a pale closed indu- mentum, upper surface puberulous. Peduncle rather stout, -+- remote from the petiole, 11/2-8 cm long, apex forked, + pseudo-umbellate, rich-flow- ered, pedicels in fr. apparently enlarging. Flowers velvety, light-brown, yellow inside, stamens yellow (ex coll.). Only seen @ buds, these depressed-glo- bose. Sepals tomentose outside. Petals pubescent outside, apex imbricating, basal parts free, blunt, rather roundish, pale green in bud apparently smaller than in A. callosa. Anthers numerous + 1mm long, on !/2-3/4 mm long filaments, sterile hardly dehiscing. Ovary depressed-globose, densely pilose, 11/2 mm high. Styles co, + 2 mm long, slen- der-clavate, overtopping flatly the anther clump. Berry acorn-shaped, 3-4 by 2 cm, brown, spotted pale. Seeds broad-elliptic, + 13/4-2 by more than 1 mm. Distr. China, Indochinese Peninsula, Hong- kong, ?Formosa, Hainan, in Malaysia: Malay Peninsula, Sumatra, Borneo. Ecol. Hill forests, rather rare, ca 900-1500 m, fl. April—July. Vern. Once noted, S. Sumatra, wait boerah. Notes. There is some variability in the size of the inflor. A. formosana HAYATA probably belongs here. Expected to occur in the Philippines. Sample treatment of vol. 1, special part. MALAYSIAN See) COLLECTORS AND COLLECTIONS CYCVOPAEDiA-O FE BOTANICAL EXPLORATION@N MALAYSIA AND A GUIDE TO THE LITERATURE CONCERNED UP TO THE YEAR 1947 COMPILED BY MRS M. J. VAN STEENIS- KRUSEMAN WITH AN INTRODUCTION BY C.G. G. J. VAN STEENIS GENERAL PART Chapter 1. Introductory essay. a. b. é: . Private collections of Malaysian ie . Use of native names. History and aim of this Cyclo- paedia. The Cyclopaedia as part of Flora Malesiana. Who is a ‘collector’? plants. . Why only Phanerogams and Pte- ridophytes? Collectors and collections of fossil plants excluded. . Correction of mistakes anderrors. . Erroneously localized plants and the sources of errors. How to cor- rect them. 1. Geography. 2. Inadequately labelled plants. 3. Plant-geographical knowledge as a control for some wrongly localized. plants. 4. Interchange of labels or wrong- ly labelled plants. 5. Intentional falsifying of labels. 6. Malaysian botanical collec- tions in which errors occur. . List of works principally con- taining illustrations of Malaysian plants, and of existing collections of drawings and photographs. . Annotated list of literature for the use of botanists and explorers in Malaysia. Nomenclature of altitudinal zones. m. Acknowledgements. Chapter 2. The technique of plant col- lecting and preservation in the tropics. Chapter 3. Chronology of the collec- tions, being a key to the history of botanical exploration of Malaysia. a. Survey of the islands separately. b. Voyages and expeditions chrono- logically. c. Early explorers up to 1840. Chapter 4. Desiderata for future explo- ration. Chapter 5. Sources of information used in compiling the special part. a. Survey of sources giving data on collectors, collections, and travels (geographically arranged). b. Reports, papers, and other infor- mation pertaining to herbaria where Malaysian collections are deposited. c. Select list of originally private collections and their present loca- tion. SPECIAL PART Abbreviations, terms, and symbols used. Alphabetical list of the collectors. (Cy- clopaedia proper, containing over 3000 names). Subject Index. NB. A geographical map and appr. 160 photographs of collectors will be in- serted. The volume will comprise appr. 600 printed pages. Aars FLORA MALESIANA [ser. I Aars, Ch. Post-Holder in Roti (Lesser Sunda Islands), sent some plants from there to Hort. Bog. in 1905 and 1917. Aart, Johannes Hendrikus van a resident of Ambon, in 1885 sent a collection of dried Sideroxylon species to Herb. Bog. The species were thought identical with those described in the work of Rumpuius. Also many living orchids and myrmecophilous plants to Hort. Bog. (1885). Aarts, F. W. J. in 1915 collected at Bodjong Terong Estate, Sidodjaja, in Priangan Residency, W. Java; 117 nos in Herb. Bog. Abaca, Y., cf. sub Forestry Bureau, Manila. Abar bin Adan, cf. sub Forest Research Institute, Buitenzorg. Abas Gelar St. Saidi, cf. sub ditto. Abbott, Dr William Louis (1860, Philadelphia, U.S.A.; 1936, Northeast, Md, U.S.A.), was educated at the University of Pennsylvania, taking his medical degree in 1884. He continued his medical studies in London. Being financially independent he decided to engage in scientific exploration and field work rather than devote his life to medical science. From 1880 on- wards he made collections of birds in America, from 1887-90 zoological collections in East Afri- ca, and subsequently visited the Seychelle Archi- pelago, Madagascar, and the Himalayas. In 1897 he explored Siam, and in the next 10 years visited Sumatra and Borneo, and cruised the coasts of Siam and the China Sea in his own vessel. He never published any of the results of his explo- rations. ITINERARY. 1901. N. Sumatra: Atjeh, at Loh Sidoh Bay, a few miles Sof Acheen Head. From May 26—-Aug. 7 islands E of Singapore, coast and rivers of Johore. From Aug.—Sept. Centr. Sumatra: Indragiri river; Lingga and Singkep.—1902. Pag(a)i Islands (part of the Mentawai Islands, W of Su- matra).—1903 and 1905. Nias, W of Sumatra. COLLECTIONS. Some plants together with C. B. Kxoss (see there) in the Pag(a)i Islands in 1902.' Living plants from Sumatra in Hort. Sing. (pres. 1903). Zoological and ethnological collections in the U.S. Nat. Mus. Washington, but no Malaysian botanical collections. LITERATURE. (1) cf. ‘Spolia Mentawaiensia’ in Kew Bull. 1926, p. 56. BIOGRAPHICAL DATA. Auk 53, 1936, p. 369— 370. Abdoelhamid (= Abdulhamid), cf. sub Forest Re- search Institute, Buitenzorg. Abdoellah (= Abdullah), cf. sub ditto. Abdoelmalik (= Abdulmalik), cf. sub ditto. 2 Abdoelmoein, cf. sub ditto. Abdoelwahab, cf. sub ditto. Abellanos(a), cf. sub Forestry Bureau, Manila. Abendanon, Eduard Cornelis (1878, Pati, Java; x), mining engineer, educated at the Technical College at Delft (Holland) and for an interval of one year at Aix-la-Chapelle (Ger- many); study tours in Europe for the D. E. Indian Government, 1900-01; in the employ of the D.E.I. Mining Service, 1901-06, from 1903-05 at his own expense making an exploration tour in China and a voyage round the world; from 1907-18 prepara- tion for, execution of, and working out of the results of the Celebes Expedition (see below) of the Royal Dutch Geographic Society;! exploration tour to Spitsbergen, 1920; Extraordinary Professor at the Municipal University of Amsterdam, 1921-25; from Nov. 1937-Novy. 1939 travelling in S. Africa, the Dutch East Indies, Australia, New Zealand and Tasmania, and the Dutch East Indies once more; from Febr. 1940—July 1946 staying at Monte Carlo; at present living at Voorburg near The Hague and planning to work out the material for a 3-volume work. He composed a geological map of the Dutch East Indies and is the author of several geological papers. Quercus him. ITINERARY. Celebes Expedition, 1909-10.‘ 1909. S. Celebes: arrival at Makassar (March 30); Ma- kassar-Palo(p)po (Apr. 10-13); Palo(p)po (13-15); reconnaissance from the N and E of the Lati- modjong Mts (15-26), e.g. exploring S. Latoepa, S. Limbong, S. Garoeang, Limpo Batoe, S. Mera- ring; Palo(p)po (Apr. 26—May 7); Ponrang (8) and exploration of the Latimodjong Mts till June 12 (top bivouac on Boeloe Palakka, May 16—June 2); ascending the middle-course of the Sa(a)dang to Rante Pao (June 14-22), also visiting Makale; Rante Pao (22-30); trip to Palo(p)po (July 1-12), e.g. visiting S. Loko; trip to the west, S. Masoepoe (8-25); along the S. Mamasa (July 26—Aug. 14), Letta Mts; descending the Lower Sa(a)dang River (17-22), Enrekang; Makassar (23-24); trip to Ma- lili (25-28); basin of the Malili River (Sept. 28— Nov. 10), visiting the lakes Matana (Oct. 3-11), Towoeti, Mahalona, Wawo toa and Masipi (Oct. 14-Nov. 3); Malili-Makassar (Nov. 10-14) and for some months back to Java.—2nd Part of the Expedition. 1910. SW. Celebes: Makassar- Palo(p)po (March 13-21); Centr. Celebes: via Lake Po(s)so to Kolone Dale (March 21—Apr. 10), visit- ing Masamba; stay at Kolone Dale (10-18); exploration of the connecting part with the E. pen-— insula of Centr. Celebes(19—23); Po(s)so (Apr. 24— May 6); the Po(s)so depression (May 7-25), visit- ing Tentena; Koro—Lariang trip (May 26-June 17), visiting Gintoe, Gimpoe, Bangkakoro, as far as_ Saloeponto; beach-bivouac at Saloeponto and by sea to Donggala (20-21); the depression, fossa Sarasina, of the SARAsINS (July 1-10), Paloe, Koe-_ abendanonii VAL. is named after vol. 1] Sample treatment Adduru lawi Plain; Donggala (11-18); by sea to Mamoe- djoe, Madjene and Pare Pare (arriving 21st); stay at Pare Pare (21-24); ancient beds of the Lower Sa(4)dang and the bay of Pare Pare (July 25—Aug. 1); back to Makassar and Aug. 7 sailing to Java. COLLECTIONS. Herb. Bog., valuable but very scanty material, few numbers, e.g. from Latimo- djong Mts.* During the expedition zoological col- lections were made too. See also under R. M. AMAD and J. J. Lerévre. Also living material for Hort. Bog. LITERATURE. (1) E. C. ABENDANON: ‘Onder- zoek van Centraal Celebes’ (Tijdschr. K.N.A.G. 26, 1909, p. 141-142, 464, 645-654, 800-821, 988— 995; 27, 1910, p. 79-106, 506-529, 979-1001, 1219- 1232); ‘Celebes en Halmaheira’ (/.c. 1910, p. 1149- 1172 and 1303; both with ill., maps, etc.); ‘Die Expedition der Kgl. Niederl. Geogr. Ges. nach Zentral Celebes 1909 und 1910’ (PETERM. Geogr. Mitt. 57, 1911, p. 234-238); ‘Midden-Celebes Expe- ditie. Geologische en geographische doorkrui- singen van Midden-Celebes (1909-1910) (Leiden 1915-18, 4 vols + atlas). (2) On the flora of the Latimodjong Mts cf. *‘Midden-Celebes Expeditie etc.’ /.c. vol. 1, p. 102; description of a new species cf. Icon. Bogor. 4', 1910, p. 81-82; and /.c. 43, 1912, p. 179-180. BIOGRAPHICAL DATA. Amsterdamsche Stu- denten-Almanak for 1926, p. 63-70-+ portr.; BACKER, Verkl. Woordenb., 1936. Abid, cf. sub Forest Research Institute, Buitenzorg. Ablaza, M., cf. sub Forestry Bureau, Manila. Aboe Baker, cf. sub Forest Research Institute, Buitenzorg. Aboe Hasan, cf. sub ditto. Aboe Oemar, cf. sub ditto. Aboeseno, cf. sub ditto. Abrahamson, E. E. in the years 1884-85 sent several North Borneo plants to Hort. Singapore. Mr HoLttum does not think that any records or specimens remain. Abram, cf. sub Forest Research Institute, Buiten- zorg. Abrams, J. a sergeant of the Forest Guards, and later Forest Ranger in Penang, /888—/9/0, obtained specimens for CH. Curtis (see there) in the Malay Peninsula (cf. BURKILL in Gard. Bull. Str. Settlem. 4, 1927, nos 4-5). Abu bin Talib joined the Forest Department in 1908; now retired. He collected mainly in Selangor, contributing to the C. F. (see sub Conservator of Forests) series of the Forest Service of the Malay Peninsula, e.g with ABDUL GHANI. COLLECTIONS. Herb. Kuala Lump., Sing., and Kew. Abyero, D., cf. sub Forestry Bureau, Manila. Achacoso, cf. sub ditto. Achmad Indonesian collector who collected specimens for K. Hryne (see there) in Simaloer Island W of N. Sumatra, from September /9/7—April 1920. COLLECTIONS. Herb. Bog.: 1818 nos; Herb. For. Res. Inst. Buitenzorg (with original labels); dupl. in Herb. Utrecht. Achmad, cf. sub Forest Research Institute, Buiten- zorg. Ackeringa, cf. AKKERINGA. Ackert, C. from Ziirich, presented museum objects from Sumatra to Bot. Mus. Univ. Ziirich in 1913. Acuna, cf. sub Forestry Bureau, Manila. Adam, cf. sub Forest Research Institute, Buiten- zorg. Adams, Arthur (1820, ? ; 1878, Honor Oak, Kent, England), Assistant Surgeon on H.M.S. ‘Semarang’ (itiner- ary etc. see sub Capt. E. BELCHER), author of some publications on this voyage,! and numerous zoolo- gical papers. Evidently some botanical collections were made during the voyage, but the chief object was decidedly zoological. Plants from a certain ADAMS are preserved in Herb. Imp. Gard. St Pe- tersb. (= Leningrad), no collecting locality known to us;? probably not identical. LITERATURE. (1) A. ApbaAms: ‘Notes on the Natural History of the islands’ (in BELCHER, Nar- rative etc., vol. 2). Together with others he published the ‘Zoology’ of the Voyage (London 1850). (2) cf. A. DECANDOLLE, Phytographie, 1880, p. 391. Addison, George Henry (1911, England; x), Assistant Curator in the Gardens Department Straits Settlements since 1938; after a short leave in India returned to Singapore in 1946; formerly Student Gardener at Kew. He collected herbarium specimens and local plants for cultivation in forest in the neighbourhood of Gap, Selangor-Pahang boundary (Aug. 1939) and at various times in and near Singapore. COLLECTIONS. Herb. Sing. Adduru, Marcelo made his first collection when a student in the University of the Philippines in 1917; he was ap- pointed Assistant of the Bureau of Science at Ma- nila in 1918, being a graduate of the Forest School. Ader FLORA MALESIANA (sero COLLECTING LOCALITIES. Philippines. 1917. Luzon: Cagayan Province (May-June). COLLECTIONS. Collection 1917, 279 nos, in Herb. Manila; dupl. in Herb. Arn. Arbor.; later collections numbered in the F.B. series (cf. Fo- restry Bureau, Manila), in Herb. Manila. In Herb. Field Mus. Chicago: 457 Philippine AET plants from a series specially collected for the Arnold Arboretum (pres. 1918) (apparently this is the AppuRU collection 1917); in U. S. Nat. Herb. Wash.: 127 Philip. plants. Ader, J. a resident of Garoet, who collected in 1928-30 in W. Java, e.g. at Kratjak, Mangoenredja, on G. Papandajan, at Bandjar, Telaga Bodas and on G. Galoenggoen (May 1930). COLLECTIONS. Herb. Bog.: Orchidaceae, Ba- lanophoraceae and Rafflesiaceae; also plants in Hort. Bog. Adér, J. W. H. a surveyor at Garoet, who sent orchids from W. Java (G. Tjikorai etc.) to Hort. Bog. (coll. 189/- 97). Some specimens, material of which is probably preserved in Herb. Bog., are mentioned in the papers of J. J. SMITH. Adjiz, Abd., cf. sub Forest Research Institute, Bui- tenzorg. 4 Administrateur Lokkibedrijf, cf. sub ditto. Aduviso, P., cf. sub Forestry Bureau, Manila. Aerensbergen, A. I. van (1874, Nijmegen, Gld, Holland; x), a priest, mis- sionary in Flores (Lesser Sunda Isls), 1908-10; at the seminary at Woloan (N. Celebes), 1910-16; at Manado (N. Celebes), 1917—October 1920; at Ba- tavia, 1920-21; at Bandoeng, 1921-25; at Buiten- zorg, 1925-28; at Bandoeng, 1928-32; and at Ba- tavia from 1932 up to the war. COLLECTING LOCALITIES. 1917. N. Celebes, Minahassa: Woloan. COLLECTIONS. Herb. Bog.: orchids from there. Aét (+ 1901; x), an Indonesian, in 1919 already in the employ of the Herbarium at Buitenzorg, in later years appointed ‘mantri’. He attended several expeditions; during some he independently made plant collections. ITINERARY. 1937. NE. Borneo. With Expedi- tion Mrs M. E. WALSH (see there). Environs of Sangkoelirang (Apr. 14-June 28), collecting at the following localities: Kari Orang (— prob. Sg. Kali- orang) (Apr. 14-20); Maloewi, G. Dajak, pinggir laoet (sea coast) and G. Batoe Ampar (22-24) and Kari Orang again; Pelawan besar and ketjil, G. Toda, G. Tembakan, G. Ketapang (low hills only), Babi Djolong, Daga Oenan, Sampajau, Malawai and Mangapoe; in the environs of Samarinda: Tenggarong (July 2—-5).—/938. Attending BUWALDA (see there) to Aroe & Tanimber Islands.—1939. With Expedition L. J. VAN Disk (see there) to P. Jap(p)en: Saroerai near Seroei-G. Wawah—Men- temboe (July 22); Seroei (23), Kainoei (25), Sam- béri (26), Saroerai near Seroei (27), Antam near Seroei (29). kp. Baroe (30), Wamiami (Aug. 1), Semémi (2), Randomi (3), Férérifi (4), Mariaroti (10), Wasabori (11-16), Sg. Soemboi near Seroei (17), Sg. Arompaul near Seroei (18), Kaunda (19), Wandési (21), Kamioraro (22-23), Sg. Papoma (24) Mariadai (26-29), G. Hirong (30), Mariadai (Sept. 1, 8), Seroei (9-11), Sg. Mémpérawaja (16-17), Soemberbaba (17); VAN D1JK and IDJAN (see there) to Biak, AT staying behind on account of ill health; Seroei (Sept. 22-—26).—/94]. With E. LUNDQUIST (see there) to W.and S. New Guinea. SW. New Guinea, near Babo: Tisa(May 8), Moetoeri (11-12), Anakasi(14~16), Jakati (19), Roriési (22), P. Kraka (23); Kaimana (June 3-4); Dutch S. New Guinea, near Oeta: Sg. Oemar (12), Sg. Si-éra (Djiera) (16-19), Najaja (20-22), Aria(26—28), Aén- doea (July 1-14), Japakopa (17), Patawai (20-21), Boeroe (23), Taréra (27-29); SW. New Guinea: Kaimana (Aug. 2); near Kaimana: Sg. Bianoga, kp. Aergoeni (9); S of Babo, Sjoga (10), Babo (13-23); near Babo: Agonda (25). COLLECTIONS. Herb. Bog.: from Exp. WALSH nos 1-654; from Exp. VAN D1iJK nos 1-845, 956-1000, for the greater part numbered too in the bb series of the Forest Research Institute; from Exp. Lunpauist nos 1-731. Dupl. Exp. WALSH in Herb. Brit. Mus. vol. 1] Sample treatment Ahmed bin Hassan From New Guinea Exp. LUNDQutsT he brought home living seeds for Hort. Bog. Agama, José (1889, Manila, Luzon, P. I.; x), a Filipino, at first Ranger of the Bureau of Forestry at Manila, and later Headranger of the Forestry Service in Br. N. Borneo; in 1926 appointed Deputy Asst Conser- vator of Forests. Several plants are named after him, especially by MERRILL. CoLLeEcTIONS. Herb. Manila: Philip. plants numbered in the F.B. series (cf. sub Forestry Bureau, Manila), and Br. N. Borneo plants (coll. 1917-19); the 2nd set of the Bornean plants in Herb. Sandakan; 23 dupl. Philip. in U.S. Nat. Herb. Wash. BIOGRAPHICAL DATA. BACKER, Verkl. Woor- denb., 1936. Agati, J. collected Helminthostachys zeylanica in Luzon, P. I., no 7799; material in Herb. Univ. Montreal. | Agoo, cf. sub Bureau of Science, Manila. - Agudo, cf. sub ditto. Aguilar, S., cf. sub ditto. Agullana in 1926 appointed Junior Ranger in the For. Dept Br.N. Borneo, collected at Sandakan (Br. N. Borneo) for the Bur. Sci. at Manila. COLLECTIONS. Herb. Manila; dupl. in Herb. Bog. (pres. 1929). Ahern, George Patrick (1859, New York City, U.S.A.; 1942, Washington, D.C., U.S.A.), Lieutenant-Colonel in the U.S. Army, organizer and Chief of the Philippine Bureau of Forestry, 1900-15.! Under his direction material of tree species was collected. In 1910 he founded the Forest School in the Philippines. After his return to the U.S.A. on duty at the Army War College, living in Washington. Canarium ahernianus MERR. was named after him. COLLECTIONS. > 850 nos collected under his direction (not by himself) in Luzon, Mindanao, etc. in 1901-02 in Herb. Manila;* dupl. in Herb. Bog. (600), Herb. Leyden, U.S. Nat. Herb. Washington (> 1800 specim.), in Berl. (210), Herb. N.Y. Bot. Garden. They were probably partly collected by QUADRAS (see there). AHERN’S collector (= RAMos, see there) numbered in the F.B. series (cf. sub Fo- restry Bureau, Manila); material in Herb. Manila too. LITERATURE.(1)G.P. AHERN: ‘Compilation of notes on the most important timber tree species of the Philippine Islands’ (1901, p. 1-112, pl. 1-43). (2) E. D. MERRILL: ‘Plantae Ahernianae’ (Dept of the Interior, Forestry Bur. Bull. 1, 1903, p.9-55). J. PERKINS: ‘Enumeration of some recently col- lected plants of AHERN, efc.’ (Fragm. FI. Philip. 1904, p. 4-66, 77-202). BIOGRAPHICAL DATA. BACKER, Verkl. Woor- denb., 1936; portr. in Philip. Journ. Forestry 2, 1939, pl. 1; Amer. Forests 48, 1942, p. 276 + portr. Ahern’s collector, cf. sub AHERN and Forestry Bu- reau, Manila. Patera AHERN Ahmad, cf. sub Forest Research Institute, Buiten- zorg. Ahmad bin A. Bakar joined the Forest Department of the Malay Peninsula in 1910; now retired. COLLECTIONS. He collected mainly in Pahang East, numbering in the C.F. (see sub Conservator of Forests) series; Herb. Kuala Lump. Ahmad bin Hassan, cf. AHMED BIN HASSAN. Ahmed bin Hadji Omar plant collector of the Singapore Botanic Gar- dens, collected in Singapore Island (cf. BURKILL in Gard. Bull. Str. Settlem. 4, 1927, nos 4-5). COLLECTIONS. Herb. Sing. Ahmed bin Hassan brother of SAPPAN BIN HASSAN and SAPPI BIN HASSAN; employed by the Botanic Gardens, Singa- pore, 1901 up to the present; 1901-12 plant col- lector to Mr RipLey; then Record Keeper, Bo- tanic Gardens. Ajat FLORA MALESIANA [ser: I COLLECTING LOCALITIES. Malay Peninsula: e.g. on Lenggong limestone cliffs;! in Singapore Island at various times. COLLECTIONS. Earlier collections numbered along with RIDLEY’s, and later in BURKILL’s Field Number Series (S.F. nos) (cf. BURKILL in Gard. Bull. Str. Settlem. 4, 1927, nos 4-5); Herb. Sing., also in Herb. of cultivated plants. LITERATURE. (1) cf. Journ. Str. Br. Roy. As. SOG470) Di). Do): - vg eA pee Be oan ? Be ot fee AJOEB Ajat, F., cf. sub Forest Research Institute, Bui- tenzorg. Ajoeb (c. 1877; x), an Indonesian, employed by the Botanic Gardens at Buitenzorg, finally as assistant ‘mantri’. He was attached to several expeditions as a plant collector. Dendrobium ajoebii J.J.S. was named after him. ITINERARY. 1910-12. Dutch N. New Guinea. With GJELLERUuP (see there).— 1914-15. Dutch N. New Guinea. With JANowskKI (see there), and later with FEUILLETAU DE BRUYN (see there).—19/6. S. Sumatra, Benkoelen. With JAcoBSON (see there).— 1920. W. Java. Garoet and environs, e.g. G. Mandalagiri and G. Djaja with LAmM.— 1920-21. Dutch N. New Guinea. With} Lam (see there). COLLECTIONS. Herb. Bog., e.g. 550 nos Ben- koelen Exp.; and especially hundreds of living plants for Hort. Bog.! 6 LITERATURE. (1) cf. list Exp. GJELLERUP in Versl. Pl. Tuin Buitenzorg for 1912, p. 22. BIOGRAPHICAL DATA. BACKER, Verkl. Woor- denb., 1936. Akker, van den collected Alternanthera sessilis R. BR. at Kesisat, Bali (Lesser Sunda Islands) in 19/4; the specimen preserved in Herb. Bog. Akkeringa, Johannes Evert (1829, Delfshaven, Z.H., Holland; 1864, Banka, E of Sumatra), engineer employed in the tinmines in Banka from about 1855; inventor of the so- called Banka drill, and author of a report on the prospects of the Billiton exploration in general and of those of the tin loads in particular.! From 1862-63 stationed in the Office at Buiten- zorg; in 1863-64 making an investigation into the occurrence of tin ore in the Riouw-Lingga Archi- pelago, in charge of the D.E. Indian government. He is said to have died from typhoid fever caught during a Borneo trip. He is commemorated in some plant names by TEYSMANN & BINNENDIK. COLLECTIONS. He sent plants from Banka to Hort. Bog.; some are described by TEYSMANN & BINNENDUJJK in the ‘Nat. Tijdschr. N.I.’? LITERATURE. (l1)cef. ‘Billiton1852-1927’ (s-Gra- venhage, M. NIJHOFF) vol. 1, p. 126. (2) cf. also SCHEFFER in ‘Observationes phyto- graphicae’ (Nat. Tijdschr. N.I. 31, 1870, p. 361, 362). BIOGRAPHICAL DATA. Nat. Tijdschr. N.I. 27, 1864, p. 31-32; portr. and some particulars in ‘Billiton 1852-1927 /.c. p. 126-127; BACKER, Verkl. Woordenb., 1936. Alambra, cf. sub Forestry Bureau, Manila. Alamsjah, cf. sub Forest Research Institute, Bui- tenzorg. Albers, E. collected near Deli Langkat in the Butta Mts, Sumatra East Coast (cf. Pflanzenreich Heft 92, p. 19). Material in Herb. Berl. Albertis, Count Luigi Maria d’ (1841, Voltri, W of Genoa, Italy; 1901, Sassari, NW. Corsica), zoologist-ethnographer who ac- companied BEcCARI (see there) on his first New Guinea trip and who afterwards made further explorations, partly alone, partly with others. In 1875 he settled in Yule Island; he made veritable raids among the natives to enrich his collections. At his arrival in Thursday Island on Jan. 4th, 1878, he was charged with murder on two China- men; after acquittal on May 4th he left Sydney, homeward bound. The genus Albertisia BEcc. was named after him and several other plant species by F. VON MUELLER and BECCARI. ITINERARY.! Jst Voyage. Sailing with BECCARI (see there) from Genoa Noy. /87/] and via Java vol. 1] Sample treatment d’Albertis (Batavia, Buitenzorg, Mt Pangrango, Soerabaja), Makassar, Timor (Dilly and Koepang) and Banda to Ambon, where preparations were made for the New Guinea trip. During the stay in Ambon, a visit was paid to Boeroe Island and to Wahai in Ceram. Sailing from Ambon (March 21, /872) via Gissar (= Geser), Ceram Laut, Goram (Apr. 1) to Dutch New Guinea (8): P. Karas (Faor), Kapour (15), Sorong (23), the 28th starting for Salawati (where fallen ill), Ramoi River (June 13), back to Sorong (July 2) and departure from there (15); coast Amberkaki (= Amberbaken) (23); P. Man- siman opposite of Doré (Aug. 3); Andai; trip in the Arfak Mts (Sept. 4-Oct. 1)? as far as Hatam; taken ill again at Andai (a month); to P. Mansiman (Nov. 2); departure to Sorong (7), arriving the 15th; Ambon (Dec. 7), sailing from there (12) in the ‘Vittor Pisani’ to Sydney, via Klein Kai (= Noehoerowa) (Doulan, Dec. 17-20), Aroe Islands (Dobo, 21st) and along the S. coast of New Guinea to Port Jackson. At the end of 1873 sailing from Australia via Fiji, Honolulu, America, to Europe, arriving in April 1874. 2nd Voyage. Nov. 10, 1874 sailing from Naples via Singapore to Somerset in Australia (stay Dec. 27-—March 4, 1875); March S sailing for Papua (SE. New Guinea): Darnley Island (12); Yule Island (arriving the 14th or 16th), which was used as a starting point for several trips:* e.g. canoe trip to the Hilda and Ethel River, Nikura and Epa (mid- April); visiting Nikura for the 2nd time (May 17) and several other trips by boat to the New Guinea coast; to Naiabui (Aug. 14), exploring the Bioto River and then going back to Naiabui (Sept. 4); march in the mountains (13-17); back in Yule Island (Sept. 22), from where Nov. 7 to Somerset on account of illness (arrival on the 14th).—/st Exploration of the Fly River* with S. MACFARLANE (see there) and H. M. CuestTEr: sailing from So- merset (Nov. 29, 1/875) in the ‘Ellengowan’; Fly River (Dec. 6-21); back at Somerset (28). — 2nd Exploration of the Fly River> with HARGRAVE and WILcox: sailing from Somerset in the ‘Neva’; Katow (May 21, /876), mouth of the Fly (22); ascending the river till June 25, travelling on foot upstream for another 5 miles and thereafter return trip during which ascending the Alice River (June 30—July 6); leaving the Fly River mouth on the 18th and via P. Mibuou(= Bristow Island) (July 18—Aug. 3), Bampton Island, Yarrou; Katow River (Aug. 7— Oct. 27, at Moatta = Mawata);7Tawan Island; back to Somerset (arrival Nov. 21).—3rd Exploration of the Fly River:® in the ‘Neva’ sailing from Somerset (May 3, 1/877), reaching the mouth on May 2Ist. As a result of d’A.’s conduct during the trip they suffered from attacks by the natives; with the Chi- nese on board difficulties arose too; in consequence they were unable to extend the journey farther inland than in 1876, only the Strickland River was detected. The return voyage was made via Moatta (= Mawata near the Katow River; Nov. 23—Dec. 5) to Thursday Island (Jan. 4, 1878). COLLECTIONS. His ornithological collection in Mus. Civ. Stor. Nat. Genoa; anthropological coll. in Mus. Florence; botanical collection, at least partly, given to BECCARI (= Florence). The plants of the trips 1875 and 1876 for the greater part described by F. VON MUELLER,’ and probably in Herb. Melbourne; those of the last trip by BEc- cARi.® According to the latter’s account the col- lection of the last exploration consists of magnifi- cent material thanks to preservation in spirits. The plants originating from the Fly River which are mentioned in d’A.’s book are numbered between 1 and 314. D’ALBERTIS LITERATURE. (1) L. M. d’Avsertis: ‘Alla Nuova Guinea. Cid che ho veduto e cid che ho fatto’ (Torino 1880, ill.); English transl. : ‘New Guinea, what I did and what I saw’ (London 1880, 2 vols); abridged French. transl. Some dates mentioned do not tally with those of BECCARI. (2) cf. ‘Una mesa fra i Papuani del Monte Ar- fak’ (Boll. Soc. Geogr. Ital. 10, 1873, p. 67-71; ‘Viaggio di de ALBERTIS nei monti Arfak e sue col- lezioni zoologiche’ (Cosmos di Guido Cora 1, 1873, p. 218-220). (3) cf. Letter from d’ALBERTIS giving some ac- count of several excursions into Southern New Guinea’ (Proc. Zool. Soc. Lond. 1875, p.530—532). (4) L. M. d’ALBerTis: ‘Remarks on the natives and products of the Fly River, New Guinea’ (Proc. Roy. Geogr. Soc. Lond. 20, 1875-76, 1876, p. 343-353, discuss. p. 353-356; extracts from letters to Dr BENNETT of Sydney, publ. in Sydney Morn- ing Herald March 1876). (5) L. M. d’Avsertis: ‘Journal of the expedition for the exploration of the Fly River in 1876’ (Syd- ney 1877) w. App. by Baron F. VON MUELLER (l.c. 7 Alberto FLORA MALESIANA [ser. I p. 14). cf. Map in PETERM. Mitt. 1878, Taf. 23; and F. ANTOINE in Oesterr. Bot. Zeitschr. 27, 1877, p. 206-208; Tour du Monde 437, 1882, p. 321-336, w. ill. (6) cf. PETERM. Mitt. 1878, p. 198-199 and 423-426; and Boll. Soc. Geogr. Ital. 15, 1878, p. V. ALDERWERELT V. ROSENBURGH 105-108; Tour du Monde 43?, 1882,p. 321-336,w. ill. (7) in ‘Descr. Not. Pap. Pl. ’’I, parts 3, 4, 5 and 6. (8) O. Beccari: ‘Notizie sulle piante raccolte dal sign. L. M. d’ALpertis alla Nuova Guinea’ (in d’ABertTis, Alla nuova Guinea, 1880, p. 571- 579); “Catalogo delle piante del fiume Fly, 1877’ (in /.c. p. 575-577), also in the Engl. transl.; ‘Sulle piante raccolte alla Nuova Guinea dal Sign. L. M. d’ALBERTIS durante l’anno 1877, con descrizione di tre nuove specie di Icacinaceae’ (in Malesia vol. 1, fasc. 3, 1878, p. 255-257). BIOGRAPHICAL DATA. Portr. in Tour du Monde 437, 1882, p. 325; Rivista Geogr. Ital. Roma 7, 1901, p. 628-632 + portr.; Boll. Soc. Geogr. Ital. 38, 1901, p. 849-855; Geogr. Journ. Lond. 18, 1901, p. 629; Deutsche Rundschau f. Geogr. u. Stat. Wien 25, 1902, p. 182-184; BACKER, Verk]. Woordenb., 1936. Alberto MERRILL cites a no 36 collected by ALBERTO, in his ‘Enum. Philip. Fl. Pl.’; no A 328, Dioscorea elmeri var. vera PRAIN & BURK., was collected at Los Banos (cf. PRAIN & BURKILL, The genus Dioscorea, 1936, p. 181); and other plants from Los Bafios, Luzon, about 1905. 8 FORTUNATE R. ALBerTO coll. for Bur. of Sci. Manila, acc. to BURKILL. Alcala, cf. sub Bureau of Science, Manila. Alderwerelt van Rosenburgh, C. R. W. K. (1863, Kedong Kebo, Poerworedjo, Java; 1936, The Hague, Holland), officer in the D.E.I. Army, 1885-1904; retired on account of deafness; tem- porary Assistant of the Herbarium at Buitenzorg, 1905-08; Conservator and later Acting Assistant of the said institution, 1910-22. Author of many systematic publications, prin- cipally on ferns. ! Dendrobium alderwereltianus J.J. S. was named after him. COLLECTIONS. Principally ferns, also orchids e.g. from Soekaboemi (W. Java) in Herb. Bog.; in Herb. Berl.: 310 nos Selaginella (pres. 1912-13). Hort. Bog.: ferns from Soekaboemi (pres. 1908). LITERATURE. (1) e.g. ‘Malayan Ferns’ (1908), “Malayan Fern Allies’ (1915) and following supple- ments. BIOGRAPHICAL DATA. Bull. Jard. Bot. Buit. sér. 3, vol. 14, 1936, p. 1-3, w. portr. a. bibliogr.; BACKER, Verkl. Woordenb., 1936; Chron. Bot. 3, 1937, p. 203 + portr. Alejandro, B., cf. sub Forestry Bureau, Manila. Alfalla, P., cf. sub ditto. Alfaro-Cardoza Portuguese. COLLECTIONS. Herb. N. Y. Bot. Gard.: 38 nos from Timor, coll. 1930. Alfiah, Moh., cf. sub Forest Research Institute, Buitenzorg. Alga, A. collected c. 20 nos in Mindanao; 6 dupl. in U.S. Nat. Herb. Wash. Ali, cf. sub Forest Research Institute, Buiten- zorg. Ali, Moh., cf. sub ditto. Ali bin Hj. Salleh joined the Forest Department in the Malay Peninsula in 1903; now retired. COLLECTIONS. Mainly in Negri Sembilan, numbered in the C.F. (see sub Conservator of Forests) series; Herb. Kuala Lump. Ali Djemar, cf. sub Forest Research Institute, Buitenzorg. Ali Silang, T., cf. sub ditto. Alimoesa, cf. sub ditto. Alkemade, J. A. van Rijn van, cf. sub RIIN VAN ALKEMADE, J. A. VAN. vol. 1] Sample treatment Altmann Alleizette, Charles d’ (1884, Paris, France; x), made botanical col- lections in Madagascar (1904-06), and in Tonkin (Indo-China, 1908). In 1909 he was appointed Underlieutenant of Administration in Oran (Afri- ca), where he was again from 1911-15 and 1917-22. He ended his military career as Chief of Adminis- tration at Versailles. CoLLEcTIONS. Herb. Paris, from Madagascar, Tonkin, etc. He collected some ferns, cult. in Hort. Sing., viz nos 516 T and 517 T in 1909 (cf. BONA- PARTE, Notes Ptéridol. fasc. 7, 1918, p. 38-39). BIOGRAPHICAL DATA. In ‘Flore générale de l’Indo-Chine’ prelim. vol. 1944, p. 33. Herb. Leyden received a set of his collection in 1948. Allen MERRILL cites the nos 168 and 171 in his ‘Enum. Philip. Fl. Pl.’, e.g. Nepenthes truncata from Min- danao. Allen, Edgar F. (born in England; x), Agricultural Officer of the Dept of Agriculture S.S. & F.M.S., stationed at Telok Anson, Perak. COLLECTIONS. Especially plants of economic interest, e.g. keladis (Araceae) in Lower Perak.! LITERATURE. (1) cf. Report Bot. Gard. Sing. for 1939, p. 2; and Gard. Bull. Str. Settlem. 11, 1941, p. 244. Allen, Reverend G. Dexter COLLECTIONS. Herb. Sarawak: a small fern from Lingga Mt., Sarawak (pres. 1906). Almagro, cf. sub Forestry Bureau, Manila. Almeida, d’ Some Garcinia leaves with 6 drawings (Garcinia hanbury Hook. f.) in Hanbury Herb. (London). Almeida, Pereira, J. d’, cf. sub PEREIRA. Almquist, Ernst Bernhard (1852, Skogs-Tibble, Uppland, Sweden; 1946, Stockholm, Sweden), Medical Officer and liche- nologist of the Swedish voyage of the ‘Vega’, 1878-79.'! He took his M.D. at Uppsala in 1882, and was professor at Stockholm since 1891. The Superintendent of botanical work during the voy- age was Dr KJELLMAN, Lecturer in Botany in the University of Uppsala. ITINERARY. Voyage in the ‘Vega’, 1878-79.! Departure from Karlskrone (Sweden) (June 22, 1878), and Gothenburg (July 4); operating in arc- tic regions, but making the homeward voyage via Japan, Hongkong, Canton; Labuan (Nov. 17-20, 1879), part of the expedition (no botanists) making an excursion to the opposite shore of Borneo; short stay at Singapore (Nov. 28—-Dec. 4); Ceylon, and home to Stockholm. COLLECTIONS. Herb. Uppsala, Herb. Stock- holm. NYLANDER described 48 lichens from Singa- pore, and 77 from Labuan.? In Labuana collection of fossil plants was made by members of the expedition; cf. also sub TREACHER. LITERATURE. (1) Baron N. A. E, NORDENS- KIOLD: ‘The voyage of the “Vega” round Asia and Europe (1878-79)’ (Engl. transl., London 1881). The official record of the expedition was edited by NoORDENSKIOLD, and consists of 5 octavo vol- umes. (2) W. NyLANDER: ‘Sertum Lichenaeae tropicae e Labuan et Singapore’ (Paris 1891). BIOGRAPHICAL DATA. WITTROCK, Icon. Bot. Berg., 1903, p. 28; I.c. 2, 1905, p. 2, t. 110, p. 206; Svenska Linné-Sallsk. Arsskr. 29, 1946, p. 66-67, + portr. Aloba, A., cf. sub Forestry Bureau, Manila. Alsacid, Godefredo L. made a study and collection of natural history specimens, e.g. of plants in Palawan (March 30- June 18, 1938); in 1939 he made two joint botanico- zoological expeditions with Carpco & EDANO to Baler and vicinity (March, May-June) in NE. Luzon. COLLECTIONS. In Herb. Manila, at least partly with B.S. numbers (cf. sub Bureau of Science, Manila). Altamirano, cf. sub Forestry Bureau, Manila. Altheer, J. J. (2; died in 1863), appointed Pharmaceutical Chemist of the D.E.I. Army in 1852 and as such stationed in Banka in 1858; for some years after 1855, Lecturer in Physics and Chemistry at the Medical School, Batavia. Author of many chemical papers in Nat. Tijd- schr. N.I. COLLECTIONS. Herb. Bog.: principally ferns;! also some Amaryllidaceae and an orchid from Banka, mentioned in TEYSMANN’S list of Banka plants;? the collection presented in 1860 to the Nat. Ver. N.I. at Batavia, was forwarded to Bui- tenzorg.* Numbering > 42; some plants numbered in the H.B. series of the Buitenzorg Herbarium. LITERATURE. (1) cf. J. AMANN (= S. KURZ) in Nat. Tijdschr. N.I. 23, 1861, p. 399-412. (2) cf. Nat. Tijdschr. N.I. 27, 1864, p. 157-258. (3) cf. Lc. 21, 1860, p. 435. BIOGRAPHICAL DATA. Tijdschr. v. N.I., N.S. 1, 1863, p. 244-245, w. bibliogr. Altmann, Hendrik (1896, Serang, Bantam, Java; Sept. 18, 1944, torpedoed in S.S. Junyo Maru when brought from Batavia to Benkoelen by the Japanese), studied tropical agriculture at Wageningen; after 6 years of practice in the Java sugar industry, appointed Group-Adviser of the Java Sugar Experiment Sta- tion at Pasoeroean; stationed successively at Koe- does (1925-31), Sidoardjo (1931-32), Pasoeroean (1932-36), Cheribon (1936-41); at the same time acting Director of the subdivision Cheribon in 1937, and Djokja (since Nov. 1941). COLLECTING LOCALITIES. 1927. Centr. Java: Koedoes. — 1930. E. Java: Modjokerto (Patjet). — 9 Alvarez FLORA MALESIANA 1931. E. Java: Soerabaja.— 1932. E. Java: Soera- baja, sf. Boedoeran, Modjokerto (Patjet, Brang- kol), Binor, Sido(h)ardjo, G. Baoeng, Gempol, Pasoeroean, Probolinggo, G. Semongkrong, G. Abang, Kepoeh, Pasangrahan Soekolilo, sf. Ta- wangsari, Tjemorolawang, Ranoe Bedali, (H)Ijang Plateau (Taman (H)Idoep), S. slope G. Smeroe.— 1933. E. Java: G. Lawoe (Sarangan and crater), S of Malang, sf. Kebonagoeng (Malang Distr.), sf. Alkmaar, Soerabaja, Modjokerto (Patjet, Bongsal), Sido(h)ardjo, Pasoeroean, Probolinggo, Oemboe- lan, G. Emprit, Tosari, sf. Goenoengsari, Ranoe Pani (G. Smeroe), Bremi, Taman (H)Idoep (Ijang plat.), Djember, Pradjekan, Besoeki, Binor, Sam- pean ravine, Idjen plateau.—/934. E. Java: Soera- baja, Porong, Trawas, sf. Krebet (Malang), Pasoe- roean, Kepoeh, Ratji, G. Bentar, (H)Ijang Plateau (Taman (H)Idoep), sf. de Maas, Besoeki, Poeger; Centr. Java: Semarang; W. Java: Buitenzorg, Ban- doeng.—1935. E. Java: G. Ardjoeno-Andjasmoro, Djoenggo, Malang Plain, Modjokerto (Patjet), Pasoeroean, G. Smeroe (Ranoe Daroengan), To- sari, sf. Pandji, sf. Pradjekan, Djember, Pasirian Poeger, Watoe-Oeloe, Besoeki, sf. Asembagoes. — 1936. E. Java: Djoenggo, Soemberbrantas, G. Ardjoeno, Pasoeroean, Kraton, Modjokerto (Pa- tjet), Poeger, Watangan, Besoeki, Garahan, Ken- deng road, G. Loeroes, Djember.—/937. W. Java: Cheribon.—/938—41. Centr. Java: in Indramajoe, Tegal and the Kromong Mts.! COLLECTIONS. Private herb. > 500 nos; many dupl. in Herb. Bog. and in Herb. Pasoeroean (453). The nos 13-58 not present in Herb. Bog. His private herbarium was taken by the Japanese, and was possibly transferred with his office to the Klaten Estate Co. W. A. TERWoGT, at Djokja. LITERATURE. (1) H. ALTMANN is the author of the botanical section in F. U. M. BUNING: ‘Het Kromonggebergte’ (Cheribon 1940). Alvarez, Ramon J. Forester of the Bur. of Forestry at Manila, since 1920 Asst Chief. COLLECTIONS. Herb. Manila, numbered in the F.B. series (see sub Forestry Bureau, Manila), etc.; 66 dupl. in U. S. Nat. Herb. Wash.; also dupl. in Edinburgh. Alviar, cf. sub Forestry Bureau, Manila. Alvins, M. V. collector employed in the Forest Dept of the Str. Settlem., from 1884-88 in Malacca. COLLECTING LOCALITIES. Malay Penin- sula.'! 1884. Reserves of Sg. Udang and Merlimau, around Selandar, Bt Danan, Naning and environs, G. Tampin, State of Sg. Ujong, E. Malacca, Ayer Kuning (from the Malacca side).— /885. Sent to Seremban (Negri Sembilan), passing through Ran- tau; Bt Lasing, Bt Sutu, Beranang, Pantai, G. Beremban; Cape Rachado (probably from Malacca by sea). Unnumbered plants (without date), bear names of Bukit Bruang, Pulau Nangka, Pulau Dodol, and other places near Malacca town, and Bt Panchor. 10 [ser 1 COLLECTIONS. His specimens (c. 1000 in 1884, 1840 in 1885 are in Herb. Sing. and have been quoted as ‘CANTLEY’s Collector’ or briefly as “CANTLEY’ (see there). He numbered in the field. LITERATURE. (1) cf. BURKILL in Gard. Bull. Str. Settlem. 4, 1927, nos 4-5. Alwi, cf. sub Forest Research Institute, Buitenzorg. Amad, Raden Mas surgeon, in 1/909 attached to ABENDANON’S Centr. Celebes Expedition;! at the beginning engaged in collecting plants, assisted by sergeant J. J. LEFEVRE; since May 20th till his departure he had to contend with illness. ITINERARY. cf. sub ABENDANON (also for liter- ature). COLLECTIONS. Herb. Bog.: some waterplants from the lakes Matana and Towoeti about Oct. 1909. He discovered the new endemic plant Boottia mesenterium HALL. f.* LITERATURE. (1) cf. E. C. ABENDANON: ‘“Mid- den-Celebes Expeditie’ vol. 1, 1915, p. 101. (2) cf. Meded. ’s Rijks Herb. Leiden no 26, 1915, Dini Amand, J. resident at Blitar (Kediri), E. Java, sent plants to Hort. Bog. in 1875. In the lists of the material preserved in Herb. Berl. acollection of mosses from Banka is mentioned (presented by J. AMAND a. 1858), this is apparently based on a mistake and should be J. AMANN (= S. KuRZzZ, see there); the same holds true for collections in Herb. Leyden, and Utrecht. Amann, J., (cf. Nat. Tijdschr. N.I. 27, 1864, p. 15 in notam, p. 403) = S. Kurz (see there). Amarillas, cf. sub Forestry Bureau, Manila. Amat, cf. sub Forest Research Institute, Buiten- zorg. Amdjah (} 1915), an Indonesian employed by the Her- barium at Buitenzorg, finally as ‘mantri’; he was attached to some Borneo Expeditions. ITINERARY. 1898-99. Borneo with A. W. NIEUWENHUIS (see there).—/909. Isl. Noesa Kam- bangan(S of Java) (June 9—20).—1912. NE. Borneo with Capt. P. VAN GENDEREN SrTort (see there for extensive itinerary of the 1st part of the trip, and for literature): arrival at Bandjermasin (May 13), starting June Sth from Pladjoe to Moeara Tagel; after breaking bivouac there, during the return voyage to Labang (Ma Sedalir), the prahu of the Indonesian collectors capsized and as result the collections and collecting necessaries were lost; in order to complete their outfit again, July Sth AMDJAH and some others went to Tg Seilor; after the evacuation of VAN GENDEREN SToRT part of the expedition started for Agisan (via the Séboekoe) by land, including AmMpJAH who went as far as Djempanga and who was back at Tenampak on vol. 1] Sample treatment Andel Sept. 27th; by prahu to Kw. Agisan (via Kapa- koean) (Oct. 4-6). Some larger explorations were made, e.g. the Tawanan Expedition (Oct. 17- Nov. 3); to Upper-Seboeda (Nov. 13), Kw. Naoen- doeng (17) and back at Tenampak (25); Dec. 7 start of the land expedition to the Simengaris; re- AMDJAH connaisance along the coast (27) and to Tawao. In the Report of the Expedition is not recorded whether the Indonesian collectors participated in these trips. The following collecting localities are mentioned (in list herbarium): B. Oeloe Se- boekoe, B.S. Tampilan, Tenampak, B.S. Toelit, G. Djempanga, Soedjau, Tikoeng, Pembliangan and Samenggaris. At the beginning of 1913 back to Tg Seilor and the greater part of the expedition leaving Bandjermasin on Jan. 10th.— 19/3. Prinseneiland (= P. Panaitan) (W of Java) (Oct.) with J. C. KONINGSBERGER; also paying a visit to the light- house Oedjoeng Koelon (W. Java). COLLECTIONS. Herb. Bog.: Borneo (1898-99) 502 nos, Java, Noesa Kambangan 242 nos, Borneo (1912) 1097 nos; Herb. Leyden: Borneo dupl. (1912). Living plants from Borneo in Hort. Bog. (coll. 1912). Amerom, Willem Frederik Hendrik van (1899, Soerabaja, Java; 1944, prisoner camp in Kiushu, Japan), Forest Officer, educated at Wage- ningen Agricultural College, went to Java in 1928. He was successively stationed at numerous places in that island; in 1934 at Garoet, and since 1935 at Madioen. Since 1940 he was attached to the Forest School at Madioen. COLLECTING LOCALITIES. W. Java: Garoet _ (1934); E. Java: Magetan (1935), Ngawi (1937), G. Lawoe (1940-41). COLLECTIONS. In Hort. and Herb. Bog.: a few numbers. In Herb. For. Res. Inst. Buitenzorg some Ja nos. Amin, cf. sub Forest Research Institute, Buitenzorg. Amiroeddin, cf. sub ditto. Amorie van der Hoeven, H. A. des (1865, Macao, China; x), in 1888 Student Gar- dener in the Botanic Gardens at Buitenzorg, and in the same year changing his situation for one in the upland cultures. He was employed again by the Botanic Gardens from 1889-91, performing pioneer work in behalf of forest-tree investigation in West Java, first as assistant of KOORDERS (see there), afterwards working independently. Later he filled several high offices in agriculture. In 1946 he lived at Leyden, Holland. Dipterocarpus vanderhoeveni K. & V. was named after him. COLLECTING LOCALITIES. W. Java: princi- pally at Tjibodas (1890), at the base of G. Halimoen near Soekaboemi (May 26, /890) and at Rongga Estate near Bandoeng. COLLECTIONS. Herb. Bog., especially ferns and material of forest trees. BIOGRAPHICAL DATA. Tijdschr. Ned. Ind. 1897, p. 1003; BACKER, Verkl. Woordenb., 1936 (cf. sub hoevenianus). Anang Indonesian, for more than twenty years in the employ of the Botanic Gardens at Buitenzorg, at present ‘mantri’. He attended some expeditions (see below). COLLECTING LOCALITIES. 1938. Moluccan expedition G. A. L. DE HAAN (see there). Collecting in Ternate: Kei Doekoe (Peak of Ternate) (Febr. 26-28); Loboso (March 5); Kabora (7); Morotai: Wajaboela (March 11-13); Goegoeti (15); G. Mo- koe (16-18); G. Toetoehoe (March 23-—Apr. 3); Halmaheira: Weda (Apr. 10-11); Tilope (13-17); Weda (May 17-24); Kobe (31).—1939. Dutch New Guinea Company Expedition (cf. sub A. SCHWARTZ). Dutch N. New Guinea: southern part of the subdi- vision Hollandia (Nimboran), basin of the Korimi River. COLLECTIONS. Herb. Bog.: Ternate nos 1-147, Morotai nos 148-393 and Halmaheira nos 394-677 from Exp. DE HAAN, New Guinea duplicates (pres. by the F.R.I.). Herb. Res. Inst. Buitenzorg: For. 106 bb nos New Guinea. Living plants in Hort. Bog. Anang Atjil, cf. sub Forest Research Institute, Bui- tenzorg. Anang Kaderi, cf. sub ditto. Andel, W. J. D. van (1867, Rotterdam, Z.H., Holland; x), came to the D.E.I. in 1891; since 1910 appointed Admini- 11 Anderson FLORA MALESIANA [ser. I strator of the Civil Service, stationed successively in the Lampong Districts (S. Sumatra) 1910-13, in Manado Residency (N. Celebes) 1913-21, in Government Celebes and Dependencies 1922-28, and in Banka and Dependencies 1928-29. After expiry of his European leave, he retired. COLLECTING LOCALITIES. J898-1902. W. Java: Soekaboemi and Djampang Tengah.—J9/2. S. Sumatra, Lampong Districts.'— 1916. N. Ce- lebes: Paleleh. COLLECTIONS. For the greater part living plants in Hort. Bog.; also in Herb. Bog., e.g. Bur- mannia lutescens from the Lampong Distr.” LITERATURE. (1) cf. Versl. Pl. Tuin Buitenzorg for 1912, p. 18. (2) cf. JonKER, Monogr. Burmanniaceae, 1938, p. 150. Anderson, Captain visited Ambon before 18/4 and from there brought seeds for ROXBURGH, e.g. of Wendlandia paniculata.'! This might be the Capt. ANDERSON of the Hon. Comp.’s ship Admiral Hughes, which called at Penang in 1790.7 LITERATURE. (1) cf. COWANin Bull. Jard. Bot. Buit. sér. 3, vol. 14, 1936, p. 33. (2) cf. TH. Forrest: ‘A voyage from Calcutta to the Mergui Archipelago etc.’ (London 1792) 10); Za Anderson, James Webster an Assistant Curator in the Gardens Department Straits Settlements, 1910-17; he later became a planter. He is the author of ‘Index of plants, Bota- nic Garden, Singapore’ (1912). After return in Bri- tain he became a private gardener. COLLECTING LOCALITIES. 1910-17. Malay Peninsula: Taiping Hills, Perak (1911) etc.— 1912. NW. Borneo, Sarawak (at all events in Aug.): visit- ing Mt Poiand probably ascending Mt Pensaung. ! COLLECTIONS. Small collections. In Herb. Kew: plants from the Malay Peninsula (pres. 1912); Herb. Sing. He numbered in the field; the speci- mens have been named and the duplicates distrib- uted in the last 12 years, but no list has been published (CorNER in litt.). LITERATURE. (1) RIDLEY in Kew Bull. 1933, p. 490 describes Microcos reticulata from Mt Pen- saung, collected by ANDERSON (without mentio- ning initials). BIOGRAPHICAL DATA. BURKILL in Bull. Str. Settlements 4, 1927, nos 4-5. Gard. Anderson, Thomas (1832, Edinburgh, Scotland; 1870, Edinburgh, Scotland), asurgeon under the Government of India and from 1860-68 Superintendent of the Royal Bo- tanic Gardens, Calcutta.! He was also the first Conservator of Forests in Bengal. In 1861 he made a journey to Java in connection with Cinchona cul- ture. In 1868 he returned to Scotland on account of illness and spent his time in working on Acan- thaceae. Eranthemum andersoni MASTERS was named after him. 12 COLLECTING LOCALITIES. 1861. W. Java, Priangan Res.: G. Goentoer and G. Malabar (Oct. 19), etc.; Buitenzorg (Nov. 6); Malay Peninsula: Singapore.” COLLECTIONS.*® Herb. Calcutta; probably > 470 nos of above-mentioned trip; Herb. Kew: In- dia, Suez (pres. 1861-70); Herb. Brit. Mus.: Sin- gapore plants; also dupl. in Herb. Berlin and Herb. Leyden. LITERATURE. (1) Author of the ‘Florula Ade- nensis’ (Journ. Linn. Soc. Lond. Bot. 1860, suppl. 1, 47 pp.). (2) T. ANDERSON: ‘On a new genus of Moraceae from Sumatra and Singapore’ (Journ. Linn. Soc. Bot. 8, 1865, p. 167-168). (3) Plants mentioned by VALCKENIER SURINGAR in ‘Het geslacht Cyperus’ (Leeuwarden 1898) p. 110; in Pflanzenreich 46, p. 271; in Bull. Jard. Bot. Buit. sér. 2, vol. 9, 1913, p. 38; Ann. Conserv. Jard. Bot. Genéve 21, 1920, p. 272. BIOGRAPHICAL DATA. Journ. Bot. 1870, p. 368 w. bibliogr.; Gard. Chron. 1870, p. 1478; PRITZEL, Thes. Lit. Bot., 1872; Biogr. Index BRITTEN & BOULGER in Journ. Bot. 26, 1888, p. 54 and in 2nd ed. by RENDLE, 1931; Curtis’ Bot. Mag. Dedic. 1827-1927, p. 151-152-+ portr.; BACKER, Verkl. Woordenb., 1936. Anderson, William (died June 3, 1778, off Anderson’s Island), Sur- geon-Botanist of Coox’s 2nd and 3rd voyage (resp. 1772-75 and 1776-80, cf. sub Cook). The 2nd voyage can be neglected, the territory treated here not being visited; on the 3rd voyage ANDERSON undertook the botany department together with NELSON (see there), but died in 1778 before arrival in the Malaysian region. He is commemorated in the genus Andersonia R. Br. COLLECTIONS. In Herb. BANKS = Brit. Mus., also MSS; no plants from Malaysia. BIOGRAPHICAL DATA. Biogr. Index BRITTEN & BOULGER in Journ. Bot. 36, 1898, p. 100, and in 2nd ed. by RENDLE, 1931; Journ. Roy. Soc. Tasmania 1909, p. 3; Journ. Bot. 54, 1916, p. 345,and /.c. 55, 1917, p. 54; Journ. & Proc. Roy. Soc. N.S.W. 55, 1921, p. 150. Anderson, William is mentioned by KRANZLIN! as collector of New Guinea plants in 1893; data on collecting localities are omitted. F. M. BatLey describes Dendrobium andersoniana, named after WILLIAM ANDERSON who collected the specimen in British New Guinea.” A certain ANDERSON (initials not mentioned) was a settler and storekeeper at Dedele (Cloudy Bay) (} 1899); he was a native of Norway, owned coco- nut plantations and began to exploit rubber, living 15 years in the colony.? He might be identical with the above-mentioned collector. LITERATURE. (1) ‘Orch. Papuan.’ in Oesterr. bot. Zeitschr. 44, 1894, p. 161-162. (2) cf. Queens]. Agr. Journ. 1901, p. 411-412. (3) cf. Ann. Rep. Br. N. G. for 1898-99, Victoria 1900, p. 13. vol. 1] Sample treatment Andrews Andersson, G. G. evidently a plant collector of H. O. ForBEs (see there). Andersson, John WICHMANN! mentions that a certain JOHN AN- DERSSON has pointed to the occurrence of ‘peat’ in Sumatra already in the year 1794. It seems probable that A. had samples, and possibly also plants from Sumatra. LITERATURE. (1) cf. Versl. gew. verg. Wis- & Natuurk. Afd. Kon. Akad. Wet. Amsterdam 29 Mei, 1909, vol. 18, 1909, p. (5)-(9). Andersson, Nils Johan (1821, Gardserum, Smaland, Sweden; 1880, Stockholm, Sweden), finished his studies at Upp- sala, taking his Ph. Dr’s degree in 1845; he was a member of a Swedish expedition round the world. In 1852 he was appointed Natural History Lecturer at Stockholm, in 1855 ditto at Lund; in 1856 Director of the Bot. Sect. of the Nat. Hist. Museum Stockholm, at the same time Professor. Author of many systematic papers e.g. on Gra- mineae and Salicaceae.! ITINERARY. Voyage in the ‘Eugenie’, 1851-53.” Departure from Karlskrona (Sept. 30, 1851) and via Madeira, Rio de Janeiro, Montevideo, Valpa- raiso, Callao, Panama, Galapagos Isls, Sandwich Isls, California, Sandwich Isls, Friendship Isls, New Holland (Sydney), Carolines, China; reaching the Philippines, Luzon: Manila (Jan. 4, 1853), mak- ing trips to Pasig, Laguna, Santa Cruz, Pagsajan, Cavinti, Louisiana, Mahayhay, Lillo, G. Banajao (not reaching the summit), Bay, Los Bafios, Lake Socal; Jan. 14th sailing for Singapore (arrival on the 25th, sailing on the 30th), where according to the account of the voyage (Dutch transl.) no col- lecting was done owing to illness; W. Java: Batavia {Febr. 4 arrival), where some little trips (Buiten- zorg etc.) were made in the last few days; sailing for the Cocos Isls on the 13th and via Keeling and Cocos Islands and the Cape of Good Hope; back in Sweden in June 1853. COLLECTIONS. Herb. Mus. Stockholm; dupl. in Herb. DECANDOLLE (Geneva) (500), St Peters- burg (= Leningrad) (1478 nos), Vienna, Liege, Brit. Mus. (mosses acquired w. Herb. HAmpe), Kew (pres. 1856-74), Berl. (128 specimens of the Gala- pagos Isl. and 55 dupl. from Stockholm pres. 1935-36), Herb. LINDEMANN (? U.S.S.R.) (59), N. Y. Bot. Gard.: 18 nos of the Philip. Isls. Evidently the labels are not very trustworthy, especially with regard to collecting localities. KUKENTHAL men- tions Cyperaceae collected by A. in Java (Batavia), Malacca, Singapore and Luzon. These data do not comply with the statement that in Singapore no collecting was done. The botany of the voyage was partly published,* but no plants of the Malaysian region were dealt with. LITERATURE. (1) The latter in DECANDOLLE, Prodromus 162, 1868, p. 190-331. (2) C. SKOGMAN: ‘Fregatten Eugenies Resa om- kring Jorden aren 1851-53 under befal af C. A. VIRGIN’ (Stockholm 1857-61, 4 vols; Ist ed. 1854~ 55) (non vidi); Dutch transl. by J.J. A. GOEVERNEUR: ‘Eene reis om de wereld met het Zweedsche oor- logsfregat Eugenie’ (Groningen 1864); German transl. by A. VON ETZEL: ‘Erdumseglung der KGnigl. Schwedischen Fregatte Eugenies in den Jahren 1851 bis 1853 etc.’ (Berlin 1856, 2 vols). cf. also some letters efc. from ANDERSSON in Svensk Vetensk. Akad. Ofvers. 10, 1853, p. 58-64, 75-76, 177-191 (in Swedish). (3) In Pflanzenreich Heft 101, IV, 20, 1936, cf. p. 71, 86, 134, 424, 519 etc. (4) Kongliga Svenska Fregatten Eugenies Resa etc. Botanik I-III, 1857-1910 by N. J. ANDERSSON and F. W. C. ARESCHOUG. Mosses by J. ANGsTROM in Ofvers. Vet.-Akad. Forh. 29, 1872-73, p. 15-29, 118-139; /.c. 33, 1876, p. 50-55; Hedwigia 14, 1875, p. 85-93. BIOGRAPHICAL DATA. Mart. Flor. Bras. vol. 1, pars 1 in ‘Vitae itiner. collect. efc.’; PRITZEL, Thes. Lit. Bot., 1872; Witrrock, Icon. Bot. Berg., 19035ep3 29 e lilvandtiny/c. 2 1905s1p 53: Andoetoe, J. H., cf. sub Forest Research Institute, Buitenzorg. Andrea, A. F. captain in the mercantile marine, presented Herb. Copenhagen with a collection of plants from the Philippines in 1875. Andreas of Menggala, W. Borneo, sent orchids to Hort. Bog. in 1896. Andreas, A. C. District Officer of the Civil Service at Tegal (Centr. Java), sent plants to Hort. Bog. in 1878. Andreas, H., cf. sub Forest Research Institute, Buitenzorg. Andreas, P. P. Doctor of Law at Buitenzorg, W. Java, presented plants to Hort. Bog. in the years /873—75; when in 1883 at Bonthain (SW. Celebes), he forwarded plants (orchids etc.) to the said Gardens. Andresen, A. J. Lieutenant Colonel in the Western Division of Borneo, sent material of the gutta-percha tree to the “‘Natuurkundige Vereeniging’ at Batavia in /852. Andrews, Charles William (1866, Hampstead, England; 1924, ?), Assistant in the British Museum, Nat. Hist. Dept, who made an expedition to Christmas Island (Indian Ocean). Some plants were named after him, e.g. Panicum andrewsi RENDLE. P ITINERARY. Left England beginning of May 1897, sailing July 23 from Batavia, and staying in Christmas Island July 29, 1897—May 1898, for 10 months.! He may have visited the island again in 1901 (see below). COLLECTIONS. Herb. Brit. Mus.: 278 plants 13 Angeles FLORA MALESIANA liserul collected in Christmas Island, pres. by Sir J. MurRAY, including types of the novelties described in the monograph of the island;? 110 spec. incl. 49 cryptogams from Christmas Island (pres. 1910) and crypt. from the same (pres. 1916). In Herb. Bog. a few duplicates, e.g. of Strongylodon lucidus SEEM., according to the label collected 12—10-/901. LITERATURE. (1) C. W. ANDREws: ‘A de- scription of Christmas Island (Indian Ocean)’ (Geogr. Journ. 13, 1899, p. 17, w. map); ‘A mono- graph of Christmas Island (Indian Ocean). Physical features and Geology. With descriptions of the Fauna and Flora by numerous contributors’ (Lon- don 1900). (2) cf. Chapter Botany in Monograph /.c. p. 171-200, pl. 17-18. BIOGRAPHICAL DATA. Who’s who 1913. Angeles, cf. sub Forestry Bureau, Manila. Angremond, Arend d’ (1883, Amsterdam, Holland; August 1945, Ja- panese prisonercamp Si Rengo-Rengo near Medan, Sumatra), was educated at the Agricultural School at Wageningen; went to Surinam as Estate Man- ager in 1905, taking much trouble for the promotion of the culture of bananas. In 1911 he went to Switzerland, where he studied for some years at Zirich under Prof. A. Ernst, taking his Ph. Dr’s degree in 1914 on a flowerbiological thesis. Subse- quently he departed for Java, where he was Di- rector of the Tobacco Experiment Station at Klaten till 1928, and later of the A.V.R.O.S. Experiment Station at Medan (Sumatra). COLLECTIONS. Probably a few plants only. In Herb. Bog.: e.g. material of Curculigo orchioides GAERTN. (pres. 193/). Angst, Ed. architect, Ziirich, presented 28 nos of cultivated plants from Java to Herb. Univers. Ziirich in 1906. Anhali, cf. sub Forest Research Institute, Buiten- zorg. Aniff = HAnirF (see there). Annandale, Thomas Nelson (1876, Edinburgh, Scotland; 1924, Calcutta, Br. India), zoologist and anthropologist, Superinten- dent of the Indian Museum, Calcutta, and Director of the Zoological Survey of India. He came to India in 1904, and is the author of many zoological papers, D. Sc. in Edinburgh 1905. He was attached to the Skeat Expedition (see below) and revisited the Malay Peninsula on more than one occasion. ITINERARY. Malay Peninsula. 1899. Skeat Expedition,'! cf. sub Yarr. ANNANDALE left the expedition after the stay at Kuala Aring towards the end of September.— /90/-—03. On some occa- sions with H. C. ROBINSON (see there). —/916. Ac- companied by a collector in the Siamese Malay States (Jan.—Febr.).” COLLECTIONS. Herb. Sing.: collection 1916, numbered in the field with S.F. nos. 14 We do not know whether any botanical collec- tions were made by him during the former expe- ditions. LITERATURE. (1) TH. N. ANNANDALE: “The Siamese Malay States’ (Scott. Geogr. Magaz. 16, 1900, p. 505-523). (2) cf. Ann. Rep. Bot. Gard. Str. Settlem. for 1916, p. 4. BIOGRAPHICAL DATA. Who’s who 1913; Rec. Ind. Mus. Calc. 27, 1925, p. 1-28, incl. bibliogr.; Journ. Bomb. Nat. Hist. Soc. 30, 1925, p. 213-214; BURKILL in Gard. Bull. Str. Settlem. 4, 1929, nos 4-5. Anonymous* Indonesia The Dutch East Indian exhibits at the exhibition at Paris in 1867 and 1900 were presented resp. in 1869 and 1901 to the Kolon. Mus. (now Ind. Inst. Amsterdam). Those of the exhibition at Amsterdam in 1882-83 are at least partly in Herb. Leyden. Anonymous Sumatra 12 Filices from Acheen (= Atjeh), N. Sumatra, with KAMEL’s plants in Herb. PETIVER in Brit. Mus.; also 3 Algae. The material must have been col- lected about 1700. A rather extensive collection from Padang and environs, Sumatra West Coast, collected in 1870, in Herb. Bog. The labels are written in a non-Ger- man hand; some of the numbers exceed 300; cf. also sub JODNER. The District Officer of the Lampong Districts (S. Sumatra) sent a lot of orchids to Hort. Bog. in 1895, The Resident of Pariaman, Sumatra West Coast, collected some plants (July 14, 1903), e.g. Knema mandaharan; in Herb. Bog. (*) Anonymous collectors are numerous in the Malaysian collections. Many of them were officials whose names can only be traced with difficulty or not at all, and who, in all probability, often did not collect in the field themselves. Some large collections were made by native col- lectors whose names are not noted (e.g. from — Borneo). Further there are quite a number of — totally anonymous collections, of which we have © not the faintest idea who made them. The ano- nymous collections cannot be neglected; some are ~ very large e.g. the ‘Native Collector(s)’ employ- ed by the Bureau of Science in Borneo. Some are — very important, and contained a lot of novelties, e.g. the grasses collected by veterinary surgeons in | Soemba Isl. (L.S.I.). Sometimes duplicates were — distributed of well-known collections with totally — inadequate labels, specially of the old collections; | these duplicates are now often ‘anonymous’. The anonymous collections have been annoying for the present compiler. They are here arranged geographically and chronologically. — vol. 1] Sample treatment Anonymous G. Meyer-Darcis, Wohlen (herbaria dealers), Switzerland, presented 408 Sumatra ferns to the Herb. Univers. Ziirich in 1905. The actual col- lector(s) is (are) not mentioned. The collection must have been made before 1905. There is a possibility that the plants are duplicates of the Sumatra collection of G. SCHNEIDER (see there). The Assistant Resident of Padang Pandjang, Sumatra West Coast, sent living orchids to Hort. Bog. in 1905. The Resident of the Lampong Districts, S. Su- matra, forwarded rattans efc. to Hort. Bog. in 1905. In Herb. Bog.: weed collection from Bah Biroeng Oeloe, Tea plantation S E of Pematang Siantar in Sumatra East Coast (coll. 1924). The Government Veterinary Surgeon of Sibolga (Res. Tapanoeli), N. Sumatra, sent more than 36 grasses from Padang Lawas to Herb. Bog. (pres. July 1926). In Herb. Kol. (= Ind.) Inst. Amsterdam: a large amount of cultivated plants, presented in /927 by the Deli Experiment Station, Medan, Sumatra East Coast. In Herb. Bog.: 16 grasses from Tapanoeli, N. Sumatra (pres. in 1927 through the intermediary of the Dir. of the Veterinary Inst.). In Herb. Kol. (= Ind.) Inst. Amsterdam: 11 wood samples + herbarium from the forests of Singkel (= Singkil, S. Atjeh, N. Sumatra) (pres. by the N.V. Houthandel Singkel in /928). The District Officer of Takengon in Atjeh, N. Sumatra, collected 4 living orchids on the Boerni Telong; in Herb. Bog. (193 .). The Forest Officer of Tapanoeli, N. Sumatra, collected 11 nos herbarium material of plants and shrubs on G. Semponan (Dairi Lands) (Nov. 27, 1929), without labels; presented to Herb. Bog. by the Forest Research Institute (Buitenzorg) towards the end of 1930. The Estate-Manager of Kotaboemi in the Lam- pong Districts, S. Sumatra, collected at least 9 nos of plants (March 22, 1934); in Herb. Bog. The Assistant Consulting Agriculturist at Wono- sarie, Palembang Res., S. Sumatra, collected the weeds nos A—M; in Herb. Bog. (pres. 1940). Many times during several years plants were sent for identification to Herb. Bog. by: 1. Deli Experiment Station, Medan. 2. General Experiment Station of the A.V.R. O.S., Medan. 3. The agriculturists in the employ of the Agri- cultural Syndicate, stationed in Sumatra West Coast and S. Sumatra. Anonymous Islands near Sumatra The ‘demang’ (native government official of the Civil Service) of the Batoe Islands, W of Sumatra, sent 2 plants to Herb. Bog. in Dec. 1930. The Civil Administrator of the said islands, ditto 1 plant to Herb. Bog. In 1/855 the Resident of Banka sent species of Nepenthes originating from there, to the ‘Natuur- kundige Vereeniging’ at Batavia; the specimens were passed to TEYSMANN, Herb. Bog., for iden- tification. The Consulting Agriculturist at Pangkalpinang, Banka, collected 4 plants (/928); in Herb. Bog. Plants originating from Billiton and P. Mendanau in Herb. Leyden, presented by the Dept of the Colonies between the years /87/—98 (cf. GODDIIN in Meded.’s Rijks Herb. Leiden no 62b, p. 20; Man- danao = Mendanau). Anonymous Malay Peninsula 12 Filices from Malacca with KAMEL’s plants in Herb. Petiver, Brit. Mus. The material must have been collected about 1700. A collector employed by THomMAs EvANs (see there) collected in P. Penang in 1808, e.g. Begonia evansiana ANDR. (cf. BACKER, Verkl. Woordenb., 1936). A Chinese collector in the employ of the Fed. Malay States Museum, made a collection of plants near the summit of G. Benom, Pahang, in the Malay Peninsula (July—Aug. 1925), at an alt. of 6000 feet and upwards. Material in Herb. Fed. Mal. Stat. Mus. = permanently on loan in Herb. Sing. (cf. M. R. HENDERSON: ‘On a collection of plants from Gunong Benom, Pahang’ in Journ. Fed. Mal. Stat. Mus. 13, 1926, p. 217-227). Anonymous Java Plants from a Javanese Garden in Herb. PLu- KENET, Herb. SLOANE 89 in Brit. Mus. The material must date from about 1700. Plants by a Dutch Gardener (from CLEYER’s garden?) from Batavia in Herb. SLOANE 286-287, in Brit. Mus. The material must date from about 1700. Java plants, collected before 1842, without the name of the collector (ex Herb. WEBB), were sold with the Herbarium of LAMBERT in 1842; they were bought by R. BROWN (= Brit. Museum). A native collector of the Botanic Gardens Bui- tenzorg, discovered Rafflesia rochussenii on the Manellawangi (= Mandelawangi, spur of G. Gedeh) on July 29, 1/850 (cf. TEYSMANN & BINNEN- DIK in Nat. Tijdschr. N.I. 1, 1850, p. 425-430, 2 pl.; and in /.c. 2, 1851, p. 651-655; cf. also HooK. Lond. Journ. Bot. & Kew Gard. Misc. 3, 1851, p. 217-220). 15 Anonymous FLORA MALESIANA [ser. I Java plants of an unknown collector in Herb. Oxford (cf. Hook. Journ. Bot. & Kew Gard. Misc. 6, 1854, p. 281). Herb. Amsterdam: Plantae Ind. Orient. Java- nicae, beautiful collection of Piperaceae etc., col- lected in the vicinity of Semarang, Centr. Java. Probably very old. The Assistant Resident of Lebak, Bantam, W. Java, sent 6 plants of Gonystylus miquelianus to Hort. Bog. in 1866. The District Officer of Rangkas Bitoeng, Ban- tam, W. Java, sent some orchids to Hort. Bog. in 1886. From Aug. /888-March 1/889 plants were collected on behalf of J. G. BOERLAGE (see there) in the environs of Buitenzorg (cf. Versl. Pl. Tuin Buitenzorg for 1888, p. 18); in Herb. Leyden. Herb. Univers. Ziirich: pharmaceutically im- portant material in spirits from Java (purch. 1895). Herb. Univers. Ziirich: 100 plants from Java, presented by Scutnz in 1903. Collector(s) unknown to compiler. An Overseer of the Forest Service collected 13 nos of plants in the mangrove forest near the Kin- derzee near Tjilatjap, Centr. Java; in Herb. Bog. (coll. 19. .). The Estate-Manager of Kiara Pajoeng, N of Tjiandjoer in W. Java, sent at least 8 nos of weeds to Herb. Bog. in July 1922. In Herb. Kol. (= Ind.) Inst. Amsterdam: material of the cultivated species of coffee from Bangelen, presented by the Malang Experiment Station in 1926. In Herb. Kol. (= Ind.) Inst. Amsterdam: samples of ‘cubeben’ + herbarium, presented by the Central Java Exp. Stat. at Salatiga in 1926 and 1931. The Consulting Horticulturist of E. Java col- lected in 1928, 30 nos Loranthaceae; in Herb. Bog. An Overseer of the Forest Service collected Rubus calycinus WALL. var. suffruticosus in E. Java, on the Jang Plateau near Taman Hidoep (Nov. 1929); in Herb. Bog. The Forest Officer of Djember, E. Java, sent 3 nos of Aleurites montana to Herb. Bog. (pres. July 1930). The Director of the Normal School at Amba- rawa, Centr. Java, collected material of Cuscuta (Jan. 1930); in Herb. Bog. 16 The Estate-Manager of Tjikopo, W. Java, col- lected Dysoxylum macrocarpum BL. (July 1933); in Herb. Bog. The Estate-Manager of Moedjoer, Pasoeroean, collected Omphalopus fallax (JACK) Naup. at Moedjoer, E. Java (cf. BAKHUIZEN V. D. BRINK in Rec. Trav. Bot. néerl. 40, 1943, p. 119); specimen in Herb. Bog. Many times during several years plants were sent for identification to Herb. Bog. by: 1. General Experiment Station for Agriculture (Algemeen Proefstation voor den Landbouw = A.P.L.) at Buitenzorg; partly collected by HACKEN- BERG, VAN HEETEREN and HUITEMA (see those) of the division Agricultural Institute, partly by others and presented through the intermediary of the said Institute; also by FRANSSEN, VAN DER GoorT and VAN DER VECHT (see those) of the division Institute for Plant Diseases (partly by others and presented by the said Institute). 2. The Private Experiment Stations, viz W. Java (a combination of the formerly separate stations for tea, rubber, efc.) at Buitenzorg, collected by the staff, principally by HEUBEL and PRILLWITZ (see those), and also a lot of material coming from various estates and presented through the inter- mediary of the Exp. Station; Salatiga, Semarang, Klaten, Malang and Besoeki (Djember) ditto, pre- sented by the respective directors. 3. Public Health Service, Medical Laboratory at Batavia, e.g. from the Malaria Department, especially waterplants. 4. The Opium Factory at Batavia. 5. The Commercial Museum (Handelsmuseum) at Batavia (the former Museum and Inquiry Office for Economic Botany, Buitenzorg). Anonymous Islands near Java The Manager of the Government Caoutchouc Estate in Noesa Kambangan (S of Java) sent some plants to Hort. Bog. in 1922. The Consulting Horticulturist of Madoera sent 2 nos of plants to Herb. Bog. in Aug. 1932. The Assistant Consulting Agriculturist at Bang- kalan (Madoera) sent some plants from experiment fields, e.g. Tenagocharis latifolia BucH. to Herb. Bog. in 1933. The Agricultural Overseer at Toendjoeng (Ma-_ doera) collected + 50 nos of weeds from experi- ment fields in 1934; in Herb. Bog. The Assistant Consulting Agriculturist at Pame- kasan (Madoera), sent respectively 21 and 13 nos of plants from experiment fields to Herb. Bog. in- 1934 and 1935. | vol. 1] Sample treatment Anonymous The Consulting Agriculturist of Madoera sent plants from experiment fields in the years 1934 and 1936; a number of them preserved in Herb. Bog. Anonymous Lesser Sunda Islands The Resident of Bali and Lombok sent a lot of living orchids to Hort. Bog. in 1886 and 1908. The Estate-Manager of Poeloekan (Bali) col- lected Salvia occidentalis SCHWARZ; in Herb. Bog. (pres. Sept. 1933). The District Officer of Lombok sent a Cypripe- dium sp. to Hort. Bog. in 1906. The Overseer of the Forestry Service collected Dysoxylum ramiflorum Mia. in Lombok; in Herb. Bog. (pres. 1933). The Veterinary Surgeon of Soembawa besar (Soembawa) sent 39 nos of grasses for identification to Herb. Bog. (1930). The Govt Veterinary Surgeon at Waingapoe (Soemba), collected Coleus scutellarioides Bru. (April 1930); in Herb. Bog. In Herb. Bog.: 5 nos of grasses from Soemba (pres. through the Veterinary Institute Buitenzorg). The Civil Administrator of Flores sent plants to Herb. Bog. in 1918. The Govt Veterinary Surgeon at Roeteng, Flores (= ? W. R. KNAApP, see there), sent 20 grasses to Herb. Bog. in 1926. A forester (in German: Forster) collected in 1932 in behalf of Herb. Berl. in Flores. cf. FORSTER, and Mrs I. RENSCH (coll. 1928). The Overseer of the Forestry Service collected prairie plants in Flores in Nov. 1934; 7 nos in Herb. Bog. (pres. Febr. 1935). The Assistant Consulting Agriculturist at Laran- toeka collected 3 nos from E. Flores in May 1935; in Herb. Bog. The Assistant Agriculturist collected 13 nos in Flores (Oct. 24, 1936); in Herb. Bog. The Consulting Agriculturist of Timor and adja- cent islands, sent plants from Larantoeka, E. Flores, to Hort. Bog. in 1938. The Civil Administrator of P. Lomblen sent a Mpristica sp. to Hort. Bog. in 1915. The Consulting Agriculturist of Timor and adja- cent islands, sent plants from P. Adonare to Hort. Bog. in 1938. The Forest Architect (=? pe Grup, see there) at Koepang, Timor, collected plants, viz in 1934 a collection of 28 nos and in April 1935: 18 nos from Fetin, Lelefoei, Nenas, Fatoe, Amnasi and Bidjeli; in Herb. Bog. The Herb. N. Y. Bot. Gard. acquired a small col- lection from Timor in 1935 (cf. Bull. N. Y. Bot. Gard. 36, 1935, p. 21). The Lieutenant of the Chinese in Ambon sent a lot of living orchids from the Tanimber Islands to Hort. Bog. in 1900. Anonymous Borneo 12 Filices from Borneo with KAMEL’s plants in Herb. PETIverR in Brit. Mus. The material must date from about 1700. The Resident of SE. Borneo sent material of some useful plants to the Nat. Ver. N.I. at Batavia in 1865; they were forwarded to Herb. Bog. (cf. Nat. Tijdschr. N.I. 29, 1867, p. 428-429, 436). The Civil Administrator of the Tidoeng lands, NE. Borneo, sent sterile specimens of Dryobalanops lanceolata BURCK to Herb. Bog. (year unknown). The Sultan of Sambas, W. Borneo, sent some iron-wood (Eusideroxylon) plants to Hort. Bog. in 1869. The Resident of Pontianak, W. Borneo, sent living plants to Hort. Bog. in 1894 and 1896; cf. sub S. W. TROMP. The Sultan of Koetei, E. Borneo, forwarded several orchids to Hort. Bog. in 1895. The Resident of SE. Borneo presented fat-yield- ing plants to Hort. Bog. in 1897. Mrs A. BONORAND, Kiisnacht, Switzerland, pre- sented fruits etc. from Borneo to Herb. Univers. Ziirich in 1902. Probably not collected by herself. In Herb. Kol. (= Ind.) Inst. Amsterdam: Pala- quium sp. div. of the Western Division of Borneo (pres. 1909). The Civil Administrator of Kotta Waringin, SW. Borneo, (=? C. VAN Nouuwuys, see there), sent living orchids to Hort. Bog. in 1912; ditto the Civil Adm. of Tanahgrogoh, E. Borneo, in 1912. A native collector has been employed by the Philippine Bureau of Science, through the agency of the Sarawak Museum during /9/3—/4. The col- lections were made in Sarawak, NW. Borneo, e.g. on Mt Merinjak, Mt Santubong, at Kuching, and Retuh (cf. E. B. CopELAND: ‘Notes on Bornean ferns’ in Philip. Journ. Sci. C. Bot. 10, 1915, p. 145-149, pl. 1). In Herb. Manila, nos ranging between 1 and 2700. Duplicates Herb. Arn. Arb., Herb. Bog., Herb. Kew, and probably elsewhere. The District Officer of W. Koetei (= ? W. C. iM Anonymous FLORA MALESIANA [serif VAN GELDER, see there), Tenggarong, E. Borneo, sent some wild orchids to Hort. Bog. in 1919. A native collector of the Sarawak Museum col- lected at Kedurong, Sarawak in 1925. Native collectors under the supervision of the Sarawak Museum collected great numbers of plants at the instigation and with the assistance, both technical and financial, of E. D. MERRILL, then at Berkeley, California. In /926—28 large col- lections were dispatched to the U.S.A. It seems probable that a duplicate set is in the Sarawak Museum. The Agricultural Officer at Pontianak, W. Bor- neo, sent specimens of Shorea to Herb. Bog. in Oct. 1929. The Consulting Agriculturist at Pontianak, W. Borneo, collected Crotalaria striata at the end of 1935; in Herb. Bog. Anonymous Philippines VIDAL in Rey. Pl. Vasc. Filip., 13, speaks of finding at the Museo del Jardin botanico, Madrid, 5 packets of Philippine plants collected in the years 1830 to 1835. In 1884 these specimens were evidently still undisturbed in their original packages. The collector is unknown, but was probably some army or naval officer. According to MERRILL (Bull. Philip. Bur. Agr. no 4, 1903 p. 33-34), CoL- MEIRO credited the collection (400 nos) erroneously to BLANCO & LLANOS. Part of it is collected on Mount Arayat in the Province of Pampanga (Luzon), in 1829; the first collection secured from one of the higher moun- tains! Anonymous Celebes The Assistant Resident of Pampanoea, SW. Celebes, sent material of orchids and Loranthaceae (dried and in spirits) to Herb. Bog. The Civil Administrator of Melillik sent a col- lection of dried plants and material in spirits to Herb. Bog. in 1912. The Assistant Resident in Boni (= Bone, SW. Celebes), sent some orchids (dried and in spirits) to Herb. Bog. in 1912. The Resident of Boni, Pampanoe(w)a, SW. Ce- lebes, sent some orchids to Hort. Bog. in 19/3. The Veterinary Surgeon at Gorontalo, N. Cele- bes, sent plants to Herb. Bog. in 1928. The Assistant Veterinary Surgeon at Donggala, Centr. W. Celebes, collected several plants in the years 1928-29; nos 1-71 originating from Lindoe, Sibalaja, Tobali, Biromaroe, etc. and nos 75-128 from Koelawi, Sidoa, Dolo and Lake Lindoe; in Herb. Bog. 18 The Assistant Veterinary Surgeon at Gorontalo, N. Celebes, collected 74 nos of plants in 1928-29; in Herb. Bog. Anonymous Islands near Celebes The Resident of Manado sent material of Gun- nera macrophylla from G. Awoe, P. Sangihe (coll. May /94]) to Herb. Bog. Anonymous Moluccas The Estate-Manager of Tobelo, Halmaheira, sent some specimens of plants to Herb. Bog. in Nov. 1919. The Resident of Ternate sent some samples of Sapotaceae to Herb. Bog. in 1884. The Post-Holder at Kairatoe, Ceram, collected at Oldenburg, by the orders of the Resident of | Ambon, specimens of Ormocarpum cochinchinense (Lour.) Merr. (July 10, /896); in Herb. Bog. (Herb. bot. var. KOORDERS no 27). A Civil Administrator of Ceram sent several specimens of sago-palms to Herb. Bog. in 1911-12. Some medicinal herbs originating from Ceram were presented by the Eykman Institute (Batavia) to Herb. Bog. in Dec. 1939. Plants from Ambon, without mentioning the collector’s name, were sold with the Herb. LAMBERT in 1842; they were bought by Ricu (cf. Advertise- ment in Athenaeum 1842, p. 44). Probably the plants were presented to LAMBERT by W. Rox- BURGH (cf. LAMBERT, Descr. of the genus Pinus, 2, 1837, appendix p. 13-24), and collected by Rox- BURGH Jr or CHRISTOPH. SMITH (see those) early in the 19th century. The Lieutenant of the Chinese in Ambon sent a lot of living orchids from that island to Hort. Bog. in 1900. The Resident of Ambon sent plants to Hort. Bog. in 1901. In Herb. Leyden: 286 Banda Plants, accompa- nied by a list of vernacular names. Neither col- lector, nor the year of collecting is mentioned, but probably dates from the first half of the 19th century. The Roman-Catholic mission in the Kai Islands sent some fodder plants to Herb. Bog. in 1927. Ditto 27 medicinal plant species to the Medical Laboratory (Batavia) in 1/939, which were for- warded to Herb. Bog. too. Anonymous New Guinea | The German New Guinea Company sent living . plants from NE. New Guinea to Hort. Bog. in 1894. The ‘Landeshauptmann’(=?SCHMIELE) informer — German New Guinea at Friedrich Wilhelmshafen ——s vol. 1] Sample treatment Arens sent living plants to Hort. Bog. (cf. Versl. Pl. Tuin Buitenzorg for 1894, p. 138, 144) in 1594. Hort. Bog.: orchids from Doré, NW. New Guinea (pres. 1899). The Resident of Ternate presented Piperaceae originating from the N. coast of New Guinea to Herb. Bog. The same in 1900 a lot of orchids from New Guinea to Herb. Bog. Herb. Univers. Ziirich: 8 New Guinea plants presented in 1904 by H. BROCKMANN, student at Winterthur. Probably not collected by him- self. In 1905 the Assistent Resident of Merauke, S. New Guinea, sent orchids to Hort. Bog. The Assistent Resident of Merauke (= prob. J. A. W. COENEN, see there), sent material pre- served in formalin from S.New Guinea to Herb. Bog. in 1912. The Officer in Command of Ambon and Ternate sent some orchids from Upper Digoel, Dutch S. New Guinea, to Hort. Bog. in 1913. In Febr. 1938, 2 specimens from Dutch S. New Guinea (Upper Digoel) were presented through the intermediary of the Army Surgeon E. M. ELSBACH at Soerabaja, to Herb. Bog. Anonymous Bismarck Archipelago The Department of Agriculture, Rabaul, New Britain (Bismarck Archipelago), sent orchids to Hort. Bog. in 1935. The New Guinea Department of Agriculture collected plants in New Britain (cf. Journ. Arn. Arbor. 22, 1941, p. 93). Ansar, S. M., cf. sub Forest Research Institute, Buitenzorg. Anta an Indonesian employee of the Herbarium at Buitenzorg, accompanied BLOEMBERGEN (see there and swb AsDAT) on his trip to Celebes and the Soela Islands in 1939, and subsequently attend- ed J. WENTHOLT (see there) on his 3rd New Guinea expedition in 1940-4] to Dutch S. New Guinea. COLLECTIONS. Herb. Bog.: about 200 nos from Merauke River. Through the outbreak of the Pacific War part of the material was retained at Makassar, especially plants collected near the Digoel River. The latter material is probably all lost. ntonio, D. collected in the Sulu Islands, cf. sub Forestry ureau, Manila. palla, P., cf. sub Forestry Bureau, Manila. Apostal, L. Forest Ranger, cf. sub Forest Department, Br. N. Borneo. Appelman, Frederik Johannes (1894, The Hague, Holland; x), studied forestry at Wageningen; in 1920 appointed Forest Officer in D. E. Indian Goyt service and as such stationed at Madioen 1920-28, in the forest district Cheribon- Tasikmalaja 1929, and since 1933 at the same time charged with the management of Garoet; in Bondowoso end of 1935—38!, in Malang 1938-40; in 1940 stationed in the Head-Office Buitenzorg for affairs connected with nature and game pro- tection; in 1941 appointed Inspector of the eastern part of the D.E.I., stationed at Makassar. After his internment by the Japanese returning thither about 1946; he retired in 1947. At present attached to the Zoological Garden in Rotterdam, Holland. COLLECTIONS. Few, Herb. Bog., e.g. from Garoet (/933), Madjalengka (1934), Rafflesia patma Bu. collected in the Penandjoeng Penin- sula? near Pangandaran on the S coast of Centr. Java (March 1934) and Rafflesia zollingeriana in S. Djember (E. Java, May 10, 1940); some Ja. nos in Herb. For. Res. Inst. Buitenzorg. LITERATURE. (1) F. J. APPELMAN: “De Baloe- ran’ (Natuur in Indié 1937, p. 49-56, w. ill.); and papers on nature protection efc. (2) F. J. APPELMAN: ‘Het schiereiland Penan- djoeng’ (Versl. N.I. Ver. t. Nat. Besch. for 1933-34, p. 55-59, fig. 3-5). BIOGRAPHICAL DATA. p. 444. Wie is dat? 1935, Arden, Stanley employed in the Agricultural Department of the Malay Peninsula, 1900-190. . COLLECTIONS. Herb. Sing. (pres. 1902) (cf. BuRKILL in Gard. Bull. Str. Settlem. 4, 1927, nos 4-5). Arendsen-Hein, Mrs COLLECTIONS. Herb. Bog.: 27 nos Brangkal, G. Ardjoeno (E. Java) in 1889. from Arens, Pedro Martin José (1884, Huancabamba, Peru; x), biologist who took his degree at Bonn (1907); employed by the Central Java Experiment Station at Salatiga 1908— 11, by the Malang Experiment Station 1911-21; subsequently Director of the Research Department of the ‘Rubber Cultuurmaatschappij Amsterdam’ (Galang, Sumatra East Coast). COLLECTING LOCALITIES. Lesser Soenda Is- lands, Bali: G. Agoeng (June 2, 19/2).—E. Java: G. Ardjoeno, Lalidjiwo (July 1, 1912, etc.); teak forest Singosari (Dec. 15, 1912); G. Kawi (Apr. 23 and Dec.9 and 12, 19/6; Jan. 30, 19/7).—Sumatra East Coast: Sg. Poetih and Prapat, etc. (/928-29); Sumatra West Coast: Alahan Pandjang (May 1930). COLLECTIONS. Phanerogams in Herb. Bog. and Herb. Leyden (Java and Bali, 150 nos); Java collections many times made together with Tu. 19 Arifin FLORA MALESIANA [ser. I WorTH (see there). Mosses, especially from later years in Sumatra, in private herbarium and Herb. VERDOORN. Arifin, cf. sub Forest Research Institute, Buiten- zorg. Aris, cf. sub ditto. Armit, Miss collected a living Dendrobium, described by Bat- LEY as Dendrobium armitae nov. spec. (cf. Queens}. Agric. Journ. 1899, p. 48), near Samarai in SE. New Guinea. Armit, William Edington de Margrat (1848, Liege, Belgium; 1901, New Guinea), Of- ficer of the Queensland mounted Police at George- town, commanded the ‘Argus Expedition’ sent by the Argus and Australasian Melbourne Newspa- pers to Papua in 1883, to report on the resources and capabilities for settlement. The main object of the expedition was to cross the southeastern penin- sula of New Guinea in ENE direction from Port Moresby to Dyke Acland Bay; on account of ill- ness and loss of one of the members they did not succeed. When the expedition was over, he made several other New Guinea trips; acting Govt Agent for Rigo and Mekeo, from July 1894-Jan. 1895 '; sub-collector of Customs, efc., at Samarai, 1895—97;. he retired towards the end of 1897; a trader of native rubber; reappointed to the public service in charge of the Northern Division during 1899. Fal- ling ill in 1900, he returned to Australia in August, sailing for New Guinea again in November when not yet recovered; finally Resident Magistrate, Northern Division. He is the author of a book and many ethnolo- gical papers on New Guinea.” The former, written over the pseudonym of J. A. LAWSON is, according to WICHMANN, wholly invented. Ficus armiti KiNG and other plants were named after him. ITINERARY. Papua, SE. New Guinea. 1883. Ar- gus Expedition:? Port Moresby (July 10); setting out for Robaduma (14), ascending a spur of the Astrolabe Range; on the way to Laloki Valley (21), Sogeri region, Meroka region at the base of Mt Belford, Aroa River; back at Port Moresby (Sept. 3).—1884. 2nd trip to E. New Guinea,‘ visiting the Moresby and Basilisk islands and the Redlick group of the Louisiades, E of New Guinea, and on the mainland: Milne Bay and East Cape. In all staying away 7 months.—/887. He is cited to have collected in this year on Mt Astrolabe (cf. sub lit. 7, BROTHERUS).—1/894. Set ashore on the NE. coast together with R. E. Guise by MAC GREGOR near Fir-tree-Point (Febr. 26), Collingwood Bay for an attempted ascent of Mt Victory:> going up Dako River (until Febr. 28), continuing by land; reaching the junction of the Tanamgina and the Waia Wai- ma (March 23, above 5200 ft); camping 10 days? near summit Mt Maneao (Mt Dayman); setting out for the return (Apr. 3), reaching the mouth of the Dako on the 12th; back on board of the ‘Mer- 20 rie England’ (14) on the way to Port Mor sby (arrival 21st). Shortly after, visit to the Goodenough Islands for exploration of the mountains.—/900. Leaving Tamata Station (Jan. 26) for Yodda Val- ley® for the purpose of discovering a practical road to the new Diggings: Ope River (Jan. 29); via Bo- rua Tutu, Tumbare Susu and the watershed of the Ope and the Kumusi; Bogi Angerita (Febr. 4); Segarata (9); the confluence of the Kumusi in the Sena (11); ascending the Kumusi to Korobama, Pidsa, Papangi; Sisureta (22), Twidi (24); Kodo on the Yodda; back at Tamata Station (Apr. 1). COLLECTIONS. Herb. Melbourne; specimens referred to by F. vON MUELLER, and others;’ 5 dupl. N. Guinea grasses in U.S. Nat. Herb. Wash. LITERATURE. (1) cf. Ann. Rep. Br. N. G. 1894/95, Brisb. 1896, p. xx. (2) ‘Wanderings in the Interior of New Guinea’ (London 1875, over the pseudonym of J. A. LAwson); ‘Notes on the Philology of the Islands adjacent to the South-Eastern Extremity of New Guinea’ (Proc. Roy. Soc. Queensl. 2, 1885, Brisb. 1886, p. 2-11; ‘The Papuans: Comparative notes on various authors, with original observation’ (in Lc. p. 78-116). (3) cf. Globus 44, 1883, p. 287; Ausland 56, 1883, p. 717 and /.c. 57, 1884, p. 255-256; Proc. Roy. Geogr. Soc. Lond. 6, 1884, p. 37-38; Boll. Soc. Geogr. Ital. 21, 1884, p. 218-225; also in the “Melbourne Argus’ 1883. (4) cf. Ausland 58, 1885, p. 480. (5) cf. Ann. Rep. Br. N. G. for 1893/94, Brisb. 1895, p. 78-87, App. 10, AA and II. (6) cf. Ll. c. 1899/1900, Brisb. 1901, p. 87-95, p. 96-98. (7) F. voN MUELLER in the Vict. Naturalist 1, 1885, p. 168; 2, 1885, p. 18-20 and 3, 1886, p. 71-72; in ‘Descr. Not. Pap. Pl.’ pt 6. A. COGNIAUx in Bull. Ac. Roy. Belg. sér. 5, vol. 14, Bruxelles 1887, p. 363. F. M. BAILEY in Queens]. Agr. Journ. 7, 1900, — p. 349. BROTHERUS described his collection of mosses in — Finska Vet. Soc. Férh. Helsingf. vols 37, 40 and 42, e.g. from Astrolabe Range /887 (no trip — known to us) and from Mt Dayman (9000 ft); cf. — also GEHEEB in Bibl. Bot. Heft 13, 1889. BIOGRAPHICAL DATA. Ann. Rep. Br. N. G. — for 1900/01, Brisb. 1902, p. xlii; MAIDEN in Journ. © Austr. Ass. Adv. Sci. Brisbane Meeting 1909, p. . 374; BACKER, Verkl. Woordenb., 1936 (1886 or 1887 erroneously stated as year of death); Journ. & Proc. Roy. Soc. N. S. W. 55, 1921, p. 150-151. Arnaud Gerkens, D. d’ COLLECTIONS. Herb. Bog.: Burmannia lutes- cens BEcc., no D 27 from Sitoehiang, S of Leuwi- liang in W. Java, collected in 1924 (cf. JONKER, Monograph Burmanniaceae, 1938, p. 151). Arnold, Joseph (1782, Beccles, SE of Norwich, England; July | 1818, Padang, Sumatra), sometimes erroneously named ARNOTH or ARNOTT; Surgeon in the British navy, 1808-16; in 1818 appointed Naturalist in the vol. 1] Sample treatment Arsin —— — ——— — service of RAFFLES who at that time was Lieuten- ant-Governor of Benkoelen (= Bencoolen). In his capacity of Naturalist he made several trips in Sumatra. Rafflesia arnoldi R. Br. was named after him and RAFELES. ! COLLECTIONS. He left behind collections of plants (acc. to BACKER, Verkl. Woordenb., shells and fossils too; the latter two were bequeathed to the Linnean Society). On his 2nd trip in S. Su- matra, he detected the mentioned Rafflesia near P. Lebar on the Manna River (2 days upstream) on May 20, /8/8. In Herb. Bog. 1 dupl., viz H.B. no 17315, originating from Siak (Sumatra East Coast). LITERATURE.(1) cf. Transact. Linn. Soc. Lond. 131, 1822, p. 201-234, r. 15-22; Flora 47, 1821, p. 637-641. BIOGRAPHICAL DATA. D. TURNER: ‘Memoir’ (Ipswich 1849); Biogr. Index BRITTEN & BOULGER in Journ. Bot. 26, 1888, p. 55, and in 2nd ed. by RENDLE, 1931; J.D. MILNER, Catalogue portraits in Kew, London 1906, p. 4; BACKER, Verkl. Woor- denb., 1936. Arnoldi, Wladimir Mitrofanowitsch (1871, Koslow, Russia; 1924, Moscow, U.S.S.R.), botanist, educated at Moscow University, where he was Assistant for several years; from 1899- 1900 he made a tour to the south of Italy, Munich and Copenhagen; in 1900 Lecturer in Moscow; in 1901 Professor at the Agricultural Institute in Nowo-Alexandria (Lublin), and in 1903-19 pro- fessor in Kharkov; subsequently working in the Kaukasus, and from 1921 in Moscow. At the outset he mainly did morphological work, but since 1909 he devoted most of his time to algological studies. With his assistant S. L. STRELIN he made a voyage to the Dutch East Indies from Jan. 10—June 5, 1909. They made a trip to the Duizend Eilanden (in the Bay of Batavia) and a voyage to the Aroe Islands to make studies on Algae.! They visited the Mariri Archipelago, E of Aroe, too. COLLECTIONS. They brought together a rich collection of demonstration material on Algae. We do not know whether phanerogams were collected too; if so, probably preserved in Kharkov. In Herb. Brit. Mus.: 7 Algae from Aroe Islands (pres. 1912); Herb. Leyden: Algae from the Malay Archip. LITERATURE. (1) cf. DAMMERMAN in Ann. Jard. Bot. Buit. 45, 1935, p. 34. W. ARNOLDI: ‘Voyage to the Malay Islands’ (Moskau 1911; in Russian) (non vidi); ‘Zur Mor- phologie einiger Dasycladaceen’ (Flora 104, 1912, p. 85-101, pl. V, 16 fig.); ‘Materialien z. Morpho- logie des Meeressiphoneen II. Bau des Thallus von Dictyosphaeria’ (/.c. 105. 1913, p. 144-161). BIOGRAPHICAL DATA. Ber. D. B. G. 42, 1924, p. (98)-(103) incl. bibliogr.; Sci. Mag. Biol. (U- kraine) 1927, p. 1-6 + portr. Arnot, D. B. (killed in Java in Febr. 1942), joined the Forest Department Malay Peninsula in Sept. 1925; In- structor of the Forest School. COLLECTING LOCALITIES, Malay Peninsula: Kanching Forest Reserve, Selangor (/927); Bubu Reserve in Perak (early part of /933) with the Forest Botanist (=? SYMINGTON); Bruas Reserve (March 1935). He collected all over the Pen- insula. COLLECTIONS. Herb. Kuala Lump.: 60 nos from G. Bubu efc., numbered in the C. F. (see sub Conservator of Forests) series. Arnoth, D. Joseph = J. ARNOLD (see there). Arnott, D. J. = J. ARNOLD (see there). Arnush, R. Andai, NW. New Guinea, sent orchids to Hort. Bog. in 1932. Arres & Ahn, are cited by BiTTer in his monograph on Lyci- anthes (Abh. Naturw. Ges. Bremen 24, 1919, p. 506), as collectors of no 3 = Lycianthes parasitica ssp. epiphytica (MERR.) Bitr. from Moeara Teweh in Borneo; this specimen in Herb. Bog. cf. VAN ASSEN & AHN, the label being misread. Arrhenius, Olof Vilhelm (1895, Stockholm, Sweden; x), physiologist-eco- logist who was educated and took his Ph. D. (1920) at Stockholm University. In Jan. 1921 he arrived at Buitenzorg in W. Java, where he worked for some time at the Foreigners’ Laboratory.' From 1920-26 Assistant at the Central Agricultural Ex- periment Station at Stockholm; on the staff of the Java Sugar Experiment Station at Pasoeroean, 1926-28; after his return to Sweden he has made researches in agriculture and forestry on his estate Kagghamra near Stockholm. COLLECTIONS. Herb. State Mus. Nat. Hist. Stockholm: material from Buitenzorg. LITERATURE. (1) cf. DAMMERMAN in Ann. Jard. Bot. Buit. 45, 1935, p. 44. Arsad, Mohammed, cf. sub Forest Research Insti- tute, Buitenzorg. Arsat retired native forest guard, cf. sub Forest De- partment, British North Borneo. Arshad an employee of the Forest Department Malay Peninsula. COLLECTIONS. In Herb. Kuala Lump., num- bered in the C. F. (see sub Conservator of Forests) series; mainly collected in Perak. Arsin (+ 1913), an Indonesian, since 1868 employed by the Botanic Gardens at Buitenzorg, finally in 1884 appointed ‘mantri’ at the Herbarium; he was an excellent connoisseur of plants, who e.g. classified the fossils of the ELBERT Expedition. In 1902 he was awarded the Silver Star for Loyalty and Merit. He arranged the Buitenzorg Herbarium in accord- ance with Index Kewensis. 21 FLORA MALESIANA [seret COLLECTING LOCALITIES. 1879. W. Java: G. Gedeh (Tjibodas, Tjibeureum, Geger Bintang) (end of Aug.—beginning of Sept.) ; G. Tjisalak near Pasir Tengah and kali Tjiapoes (beginning of Oct.); Pa- meungpeuk (Oct. 21).—/883. Sumatra West Coast: with Dr Burck (itinerary etc. see there) to the a ARSIN Padang Highlands (Aug.—Nov.).—/885-—86. E. New Guinea: with H. O. Fores (itinerary efc. see there). —1905. Krakatau in Soenda Straits: with Tu. VALETON (see there) on the Ist of March. COLLECTIONS. Herb. Bog.; Java plants num- bered in the H.B. series. The collections from Sumatra, New Guinea and Krakatau probably re- spectively under the names of BURCK, FORBES and VALETON. Asda(t) a Sundanese employed by the Herbarium at Bui- tenzorg, who attended BLOEMBERGEN (see there) on his trip to the Soela Islands in 1939, and a soil- scientific expedition under the direction of VAN DER VoortT and TANZER (see those) of the Soil Science Institute (Buitenzorg),-to the environs of Troemon in SW. Atjeh (N. Sumatra) (Aug. 17-Sept. 2, 1941). COLLECTIONS. Herb. Bog.: 207 nos Sumatra plants; Herb. For. Res. Inst. Buitenzorg: 69 nos Soela Islands, numbered in the bb. series, collected by AsDA & ANTA. Asgar, cf. sub Forest Research Institute, Buitenzorg, 22 Askey, A. M. Ranger in the Forest Department (1906-22), col- lected in the Malay Peninsula (cf. BURKILL in Gard. Bull. Str. Settlem. 4, 1927, nos 4-5). COLLECTIONS. Herb. Kuala Lump., numbered in the C: F. (see sub Conservator of Forests) series. Askey, J. E. Ranger in the Forest Department (1905-10), col- lected in the Malay Peninsula (cf. BURKILL in Gard. Bull. Str. Settlem. 4, 1927, nos 4-5). CoLLectTions. Herb. Kuala Lump., e.g. from Negri Sembilan, the Dindings, and mainly Perak and Penang; numbered in the C. F. (see sub Con- servator of Forests) series. Asloeri an Indonesian, since 1910 employed by the Ex- periment Station for the Java Sugar Industry at Pasoeroean, in 1915 appointed ‘mandoer’ of the selection division and since 1937 Head-mandoer of the gardens. He has a notorious capacity for classifying still undescribed cane clones. He as- sisted many members of the staff of the said insti- tution during trips in and oudside Java, e.g. Prof. Dr J. JeswieTt, Dr C. A. BACKER and Dr O. POsSTHUMUS (see those). COLLECTING LOCALITIES. E. Java: Pasoe- roean and neighbourhood (/93/—33); G. Kawi 1932). COLLECTIONS. Herb. Pasoer.: 47 nos. Asnawigana, cf. sub Forest Research Institute, Buitenzorg. Assen, van & Ahn collected some plants near Moeara Teweh in SE. Borneo (June 24, 1/894); specimens in Herb. Bog. Assu, Abd., cf. sub Conservator of Forests series, Kepong. Atang, cf. sub Forest Research Institute, Buitenzorg. Atasrip (7 1921), an Indonesian, since 1888 employed by the Buitenzorg Herbarium, finally ‘mantri’.! He assisted during some expeditions outside Java (see below). Dryopteris atasripii ROSENST. was named after him. ITINERARY. 1/899. Moluccas, with expedition Ham (see there). Collecting in Banda (Jan. 27), Ter- nate (Febr. 2); P. Obi (March 22—Aug. 12), and in this period visiting P. Bisa, Obi-Latoe (July), P. Belang Belang and Woi Besar (July); Ternate (Aug. 23).—1903. Dutch North New Guinea. With Expe- dition WICHMANN (see there, and sub DmBJA), near Geelvink Bay, Cyclop Mts, Lake Sentani, Hol- landia, etc. ATASRIP returned to Java before the fixed date, viz when the trip to Lake Sentani and the Cyclop Mts was over.” CoLLectTIONS. Herb. Bog.: 130 nos from the Moluccas, Banda 3, Ternate 43, Obi Isls 84; c. 250 nos from New Guinea; dupl. in Herb. Leyden. He vol. 1] Sample treatment Baalen brought home living plants from New Guinea too.” The New Guinea plants were provisionally classi- fied by ArsIN and later described by VALETON.? LITERATURE. (1) In the papers published by WICHMANN and LoreNTZ on this New Guinea expedition, he is erroneously mentioned as ATJIP or Apyip. ATASRIP was accompanied by a 2nd as- sistant, viz DsrpJA (see there). (2) cf. Versl. Pl. Tuin Buitenzorg for 1903, p. 102-103; Bull. Mij Bev. Nat. Ond. Ned. Kol. no 45, p. 6. (3) In ‘Plantae papuanae’ (Bull. Dép. Agr. Ind. néerl. 10, 1907, 72 pp.); cf. also in FeEDDE Repert. 5, 1908, p. 377-397). cf. also J. J. SmirH in Nova Guinea vol. 8. BIOGRAPHICAL DATA. BACKER, Verkl. Woor- denb., 1936. Atin, cf. sub Forest Research Institute, Buitenzorg. Atije(h) Indonesian, in the employ of the Botanic Garden Buitenzorg. ITINERARY. /9/3—14. With Expedition VAN HULSTIIN (see there) to the Soela Islands. Accord- ing to Versl. Pl. Tuin Buitenzorg for 1913, his departure from Buitenzorg took place on July 18, 1912; this is, apparently, a printer’s error and rightly should be /9/3. Collecting was done in the following islands: Soela besi (= Sanana), Mangoli (e.g. on G. Pakao), Taliaboe, P. Masonie, P. Seho and P. Kano. During the latter part of the expe- dition VAN HULSTIJN was assisted by SAANAM (see there). COLLECTIONS. Herb. Bog.: 420 nos Soela Is- lands Exp. VAN Hu stun. Living plants in Hort. Bog. Atmoesoewarno, cf. sub Forest Research Institute, Buitenzorg. Atmotaroena, cf. sub ditto. Augustin, D., cf. sub Forestry Bureau, Manila. Augustinovicz, Thoma Matveyevich a medical man and zealous botanical collector. From 1871—82 he explored E. Siberia. In /879, when on his home voyage in a Russian ship, he gathered some plants in Singapore (cf. E. BRETSCHNEIDER, History of European discoveries in China, London 1898, p. 1035 (The cited christian name Foma is an error). COLLECTIONS. In Herb. Bot. Gard. St Peters- burg (= Leningrad). Awang Lela bin Mukin Adam joined the Forest Department Malay Peninsula in 1926. ; COLLECTIONS. Herb. Kuala Lump., numbered in the C. F. (see sub Conservator of Forests) series; mainly collected in Pahang. Dupl. in Herb. Sing. Azaola, Inigo Gonzales y a planter in Lagufia Prov., Luzon, mentioned by MERRILL as cited in literature as a collector of Philippine plants (cf. Bull. no 4 Bur. of Agr. Manila, 1903, p. 30). The genus Azaola BLANCO and several other plants were named after him. He collected some Ra fflesia specimens on Mt ATIJE Majaijai (cf. HtiERONyMus, Ueber Rafflesia scha- denbergiana GOpp. efc.’ (Breslau 1885, p. 8). He sent his plants to BLANCO, a friend of his, therefore they probably do not exist any more. Aziz bin Ahmad joined the Forest Department Malay Peninsula in 1914 as Forest Ranger; retired now. CoLLectTIons. Herb. Kuala Lump., numbered in the C. F. (see sub Conservator of Forests) series; mainly collected in Pahang West. Azurin, cf. sub Forest Research Institute, Buiten- zorg. bb. nos, cf. sub Forestry Research Institute, Bui- tenzorg. Bit. nos, cf. sub ditto. B.S. nos, cf. sub Bureau of Science, Manila. Baalen, J. van collected some weeds from Tjikantjoeng Rubber Estate near Tasikmalaja (Priangan Res.), W. Java, 23 Backer FLORA MALESIANA [ser. I, vol. 1] at an altitude of + 260 m (June 24, 1922); plants in Herb. Bog. Babak, cf. sub Forest Research Institute, Buitenzorg. Bacani, E. S., cf. sub Forestry Bureau, Manila. Bachmid, Mohammed, cf. sub Forest Research In- stitute, Buitenzorg. Backer, Cornelis Andries (1874, Oudenbosch, Holland; x), a schoolmaster who came to the D.E.I. in 1901 and when stationed at Weltevreden (Batavia) at once enthusiastically started collecting and studying Java plants. He came into touch with Trev, at that time Director of the Botanic Gardens at Buitenzorg, through whose intermediary he was appointed at the Her- barium of the said institution in 1905, since 1914 Botanist for the Java flora. He spent many years in travelling in order to get the required materials for a flora of Java.' He was pensioned off at the end of 1924 and after that was temporarily employed by the Experiment Station for the Java Sugar Industry at Pasoeroean (1925-31) for composing a weed flora of the Java sugar-cane fields. When the latter book was fin- ished he left for Holland, settling at Heemstede (near Haarlem), where he up till now is engaged in continuing his work on the flora of Java which is near its completion. In 1936 he was awarded a honorary Dr’s degree at Utrecht University. He is the author ofa ‘Verklarend Woordenboek’, 1936, many times cited in this cyclopaedia. Many plants were named after him including the genus Backeria BAKH. f. (Melast.). COLLECTING LOCALITIES. 1903-05. W. Java: environs of Batavia, e.g. at Pepango, Sentiong, Pe- sing, Doeri, Kemajoran, Bidara Tjina, Mr Cornelis, Tg Priok, Angké, Antjol, Tanggerang etc.; G. Tangkoeban Prahoe (Oct. 2, 1903).—/905—24 col- lecting many times in the environs of Buitenzorg, e.g. on G. Salak (Tjiapoes, Waroeng Loa), at De- pok, Sémplak, Masing, Tjibinoeng, Tjilodong, Tjisééng, Koeripan, G. Tjibodas, Tjampea, Leu- wiliang, Bodjong Gedeh, Klappa Noenggal, Ba- toetoelis, Tjiogrék, G. Gedeh (Tjibodas), Tjianten, Bolang, Djasinga.—/906. With Ernst & PULLE:? P. Edam (Bay of Batavia) (Apr. 24), Tg Rata (= Vlakke Hoek) (Lampong Distr., S. Sumatra) (25), Java’s Eerste Punt (W. Java) (25), Krakatau & Ver- laten Eiland (26).—1908. Krakatau & Verlaten Ei- land (May 4-5), Lang Eiland (6).4A—1911-15, trav- elling all over Java.—1911. Centr. Java: Poerwa- karta, Maos (Apr. 11); Batoe Raden (12), G. Sla- mat (upwards of 1000 m) (13); Batoe Raden, S. slope of G. Slamat (16-22); W. Java: environs of Buitenzorg; in Bantam (June 9-July 1): Rangkas Bitoeng, TyjiléJés, G. Kentjana, G. Kendeng, Ma- lingping, Panjawoegan, Bajah, G. Madoer, Lang- kop, Tjitorék, Moentjang, Pasir Agoena and envi- rons, Rangkas Bitoeng; Dec. 21-30 visiting the south coast: Tjibadak, Wijnkoopsbaai, Tjisolok, Tjilétoeh, Zandbaai, Tjiképoeh; Dec. 31—-Jan 2, 24 1912: Pasawahan, Tjiémas, Palaboehan Ratoe.— 1912. W. Java: in Priangan Res. (March 1-5): en- virons of Bandoeng, Lembang, G. Tangkoeban Prahoe, Nagreh (= prob. Nagreg); trip to Djokja (= Jogjakarta) and the south coast of Centr. & E. Java (Apr. 6-19): Djokja and environs, Wonosari, BACKER Kemadang, south coast, Djepitoe, Kalak, Patjitan, Toelahan, Tegalombo, Slahoeng; E. Java (May 27-June 19); Bangil, Malang District near Wono- kerto, Soemberwalo, Gondang Legi, Ampel Ga- ding, Kali Glidik, Widodaren (near G. Smeroe), south coast, Soerabaja; W. Java: Leuwiliang & Bo- lang (July 16-19); trip to Bandjar in the SE. part of W. Java and Noesa Kambangan § of Centr. Java (Aug. 27-Sept. 11): Bandjar, Rawah Lakbok (27), G. Babakan, Tjikawoeng, Rawah Apoe, Tjikem- bo(e)lan, Tjilatjap, Noesa Kambangan (Sept. 7-8), Bandjar, Wanaredja (11); W. Java: Tjiandjoer and environs & G. Gedeh-Pangrango (Sept. 13-21); trip to environs of Cheribon (Oct. 17—31): Cheri- bon, Linggadjati, G. Tjeremai (up to 2210 m), Koeningan, G. Tjeremai (3075 m, crater); Prian- gan Res. (Noy. 12—Dec. 1): Bandoeng, Garoet- Tjipanas, G. Goentoer, Waspada, G. Tjikorai, Tjisoeroepan, G. Papandajan, Taloen, Tjinjiroean; in Krawang District (Dec. 19-Jan. 1, 1913): Tji- leungsi, Klappa Noenggal, Si Boentoe, Goea Gad- jah and surroundings, G. Handjawang, G. Boetik Boeligir, G. Karang Gantoengan, G. Soenarari. —1913. Centr. & E. Java (Jan. 21—Febr. 12): Solo, Karanganjer, Soemberlawang, Goendih, Koe- Sample treatment of vol. 2 MEAP a ANP AWN ToL 1B E BEING A MUCH ENLARGED SECOND EDITION OF MALCBISCHE VEGETATLIESCHETSEN’ ILLUSTRATED BY NUMEROUS PLANT FIGURES PHOTOGRAPHS AND MAPS BY C. G.-G.. J. VAN STEENIS Contents: Introduction to the physiognomy of tropical vegetation. Chapter 1. Physiognomical vegetation maps of Malaysia. Methods of defining vegetation types, forest analysis, aerial surveys, hints for collecting, &c. Chapter 2. General principles of dynamics in vegetation types: influence of man, seral communities, grass and forest fires, pioneer plants, afforestation, altitudinal zonation, new scheme for factors determining the final composition of the vegetation, &c. Chapter 3. Soil and flora in Malaysia: gass wells, fumaroles, inundation, guano plants, slip soils, nitrate plants, crater gases, inland halophytes, biogenous lime formation, edible earth, &c. Chapter 4. Climate and flora: macroclimate versus microclimate, strength of the dry season, difference between E. and W. slopes, effects of drought generally, in the mountains and in the swamps, frost plains, effects of lightning, &c. Chapter 5. Description of vegetation types proper: a. Open vegetations, nos 1-100. 5. Savannah vegetations, nos 101-150. c. Secondary forest types, numbers 151-200. d. Primary vegetation types. 1. Conspicuous life forms (cycads, tree ferns, palms, Pandanus, bamboos, other Monocotyledonous plants), nos 201-274. . Casuarina forests and conifers, nos 275-300. Monsoon forest types, nos 301-360. . Mangrove types, nos 361-370. . Swamp- and peat forest types, nos 371-400. . Dry-land forests. a. Mixed forests in the lowland, nos 450-600. 6. Mountain forests, nos 601-700. e. Anthropogenic vegetation, nos 701-800. DAnhRwWN N.B. The volume will comprise appr. 600 printed pages. The MS. will probably be completed within two years.—ED. Fig. 8. Pandan vegetation in the crater of Mt Kaba, SW. Sumatra. (DE Voocp) [vol. 2] Sample treatment 3 361. Dolichandrone spathacea (L. f.) K. Scu. (Bign.).—Poko kulo (Mal. Pen.), koeda koeda, m, (ki)djaran, s, (kajoe) djaran, j, (djo)djamé (Molucc.), Mangrove trumpet tree. Small to medium-sized soft-woody tree widely distributed from Malabar throughout Malaysia to New Caledonia (not known from Australia), re- stricted to the littoral zone, mostly in tidal creeks, estuaries in the back-mangrove. Leaves 1-pinnate, flowers very long, tubular, white, nocturnal, in short racemes, apparently pollinated by long- tongued moths. Pods linear falcate, seeds corky, rectangular. In Perlis, N-most prov. of the Mal. Peninsula, it is a feature of the low country, stand- ing like an upright poplar in the rice-fields, onwards from Kodiang, and flanks the roads which lead to Kangar and Singgora. Nowhere else it is reported gregarious or of physiognomic impor- tance (1). LITERATURE. (1) CORNER, Wayside trees Ma- —laya (1940) 164, Atlas pl. 26. 139. Antidesma ghaesembilla GAERTN. (Euph.).— Onjam, s, démpoel lélés, j, kénjan, j, sépat, j, koe- tikata goenoeng (Ambon), goentjak, koentsjoh (Mal. Pen.). Rather small often crooked tree 5-13 m by 10-25 cm (1) from trop. Africa and SE. Asia to West Ma- laysia, fruits edible, both native and planted (2). Crown dense, bushy, first conical then round, or umbrella-shaped, very twiggy (3). In the Philippines this is one of the commonest trees of the mixed monsoon forest and characteristic of open grass- lands. Steup found it one of the most common savannah trees in Celebes (4) dominating on the extensive hills and flats of Mamoedjoe (SW. Cel.), near Galoempang, in orchard facies. StEup found also locally dominance in the Masoepoe savan- nahs; elsewhere it occurs mixed. BLOEMBERGEN found it dominating in grass-thickets on terraces in the S of Soela Sanana Isl. at 450 m alt. It deci- dedly prefers a pronounced dry season. LITERATURE. (1) HeEyng, Nutt. Pl. N.I. (1927) 186. (2) BuRKILL, Dict. Ec. Pr. (1935) 186. (3) Cor- NER, Wayside trees (1940) 233. (4) De Trop. Natuur 25 (1936) Jub. Nr p. 43-5. 225. Pandanus L. (Pandan.).— Pandan, screw-palm, Screw-pine. Pandanus is an old Old World genus of the Pan- danaceae distributed from Africa towards SE. Asia, Malaysia, Australia, and the Pacific Islands. Its area includes Tahiti, New Caledonia, E. Australia to 33° SL, Madagascar, trop. Africa, Assam, Ton- kin, S. China, Riu Kiu and Bonin Isl. There is no marked centre of specific development. Although monographic treatments have been given by S. Kurz (1), So-ms LAUBACH (2), WARBURG (3) and later additions by MARTELLI (4) identification of species is very difficult, specific discrimination of the more than 300 species described being far from satisfactory. This induced recent students of Pa- puan specimens [KANEHIRA and MERRILL & PERRY (14)] to add to the scores of endemics. This is partly due to the fragmentary collections of these generally bulky dioecious plants. For a proper placing in the subgenera both Q and dplants are needed; the d are in many species unknown. BACKER in a careful study at Buitenzorg (5) re- duced several species described from Java. A new monograph prepared in Malaysia is urgently re- Fig. 1. P. faviger BACKER, on edge of Lake Bratan, Bali, young specimens, ca 1000 m alt. quired. Though not occurring in the temperate parts of the globe (except, possibly, as fossils) pandans grow in the mountains of New Guinea up to over 3200 m. No sandy beach is devoid of pan- dans, except in dry regions which they avoid. I gave a provisional general treatment of the role pandans play in the Malaysian vegetation in 1935 (6). The general habit of these characteristic woody plants (fig. 1-2) is rather uniform: a terete soft- woody smooth stem is supported below by con- spicuous straight stilt or prop roots; a tap-root is absent. Sometimes these roots sprout from all over the stem and a clear bole is nearly absent, and in some cases they are also found on the branches. The upper roots often do not reach the soil and remain aerial (fig. 2, 3). The stem is ringed by the scars of the sheathing amplexicaulous leaves. It is thinnest at its base. The apex is mostly pseudo- dichotomously branched with densely tufted foli- age (fig. 1) as in Dracaena (candelabra or palmid habit). The inflorescences are terminal, and cause sympodial branching. Small species are sometimes nearly stemless, e.g. P. caricosus KURZ and P. polycephalus LAMK. non SPR. measuring 3/4—11/2 m. Others, e.g. P. bidur JUNGH. from the beach and P. brachyphyllus MERR. & PERRY from the Papuan Mts may reach 20-25 m in height, while in P. jiulia- nettii MART. the average large tree is as much as 30 m tall (fig. 5). A very unusual character is found in P. aggregatus MERR. & PERRY where stems form clumps of several trees rising from a small com- pact group of erect prop roots (14, p. 177). The conspicuous stilt roots are mostly straight and un- branched, although when wounded they may branch profusely into much thinner roots. They occur both in specimens growing in dry or inun- dated soils and bear small prickly warts in longitu- dinal rows (fig. 3). In large species the root caps are larger than a fist and consist of thin corky sheets. 4 FLORA MALESIANA [ser. I Sometimes epiphytes and other plants settle in these ‘hanging flowerpots’; I illustrated this (7) from Bali (fig. 3). The seeds of these plants were probably mostly dispersed by ants. In greenhouses the roots in the soil sometimes produce very thin short negatively geotropic rootlets, as palms some- Fig. 2. Large stilt-root and aerial root system of P. faviger BACKER, mixed rain-forest, near Lake Bratan, Bali, 1000 m alt. times do in inundated places. Similar tiny upturned adventitious roots are sometimes found on the stem within the leaf sheaths (similar to palms) (8) and apparently later harden into upturned prickles (14, p. 157, 163, 165). The linear leaves of Pandanus are set in 3 spirals accentuating the characteristic stiff habit. The prickles on their margin and those on the underside of the midrib are directed up- wards; sometimes additional prickles are found on the upper side of the parallel nerves. Cultigens are sometimes unarmed. The midrib portion of the leaves is shallowly sulcate (fig. 6), and acts as a furrow along which débris assembles on the often long persistent leaf base. There is an axillary bud. The width of the leaf varies between 1 and 20cm, and in the larger species the leaves may reach a length of 4'/2m. Young foliage is mostly pale but in P. houlletii CARR. it is coppery- purple. The pine-apple-like fruit consists of the fused fruits of the individual 9 fls each topped by an indurated style. They vary considerably in size and inner structure in different sections of the genus (fig 6,9); in P. leram Kurz the separate fruits from the spadicea] capitulum may attain 12 by 8 cm; all are more or less obconical. Of P. simplex MERR., a medium-sized Philippine species, unbranched stem 6 m, ELMER collected a solitary syncarp 60 by 20 cm weighing ca 25 kg, the fruits were gradually shed from the apex to the base of the syncarp (15). In small species the capitulum measures only 2 cm. The pyrenes are surrounded by a pulpy mesocarp full of fibres; in several species the ripe fruit is a beautiful bright red, and the mesocarp is often rich in red fat (carotines) and aleurone (13). They germinate from the base. They are probably partly water- and partly animal-dis- persed. The male flowers consist of branched spa- dices with an abundance of stamens; the spadix is provided mostly with large thin cream-coloured bracts; these infl. are sometimes very fragrant (e.g. in P. tectorius PARK.), sometimes they are fetid. The method of pollination is unknown, probably by wind or partly by bats. The frequency of flow- ering in several species is often low, and the num- ber of d and Q plants is often very different. Some- times 6d, in other cases 9, plants predominate. In many species d plants are as yet unknown impeding taxonomical study. Some cultigens never flower. Most species are terrestrial, a few are epiphytic, e.g. P. epiphyticus MART. from Borneo; BECCARI mentions (9) 2 species from Sarawak associated with epiphytic ferns and orchids; in the Malay Peninsula there are also two epiphytic species pref- erably growing on trees alongside rivers. Epiphytes are sometimes found abundantly on the smooth, pandan stems (VERSTEEG in sago swamps in New Guinea, cf SMITH in Nova Guinea 8). Near Bui- tenzorg Dendrobium pandaneti RIDL. is mostly confined to Metroxylon! Brass found Hymeno- lepis validinervis KUNZE, a fern, restricted to pan- dan crowns at 3225 m at Habbema camp in W. New Guinea. Several Pandans are cultivated in Malay- sia, either for ornamental or other purposes. Two forms of unknown origin are cultivated for the fragrance of the drying leaves, viz P. amaryllidifo- Fig. 3. ‘Hanging flower pots’ from Bali. Root caps of P. faviger BACKER with epiphytic plants, mostly ferns, ca 1000 m alt. vol. 2] Sample treatment 5 lius Roxs. and P. Jatifolius Hassx. (P. odorum RIpL.) The latter cultigen is only known to be sterile, the former only as 6 individuals, both are sparsely armed. An unarmed form of P. tectorius ParK., f. Jaevis WARB., is cultivated for the fragrant bracts of the infl. Of the same species a f. samak vitamins (carotenes). Mountain tribes of Papuans save the wild trees in their clearings (fig. 4, 5) and also plant them in groves. These Papuan species may become industrially important. P. subumbel- latus Soims, P. jiulianettii MArt., P. conoideus LAMK., P. houlletii Carr., P. leram Jones, P. Fig. 4. View up lower valley of Bele river, W. Central New Guinea, from 2140 m on E slope. Old garden lands of Papuans. Newly planted sweet potato gardens on left, and others in distance. Groups of spared and planted Pandanus prob. brosimos MERR. & PERRY. Numerous planted(?) Casuarina trees on deforested slopes, and relic patches of Castanopsis forest. Dec. 1938. (BRAss, 3rd Archbold New Guinean Exp.) Wars. is a group of prickly pandans cultivated for the tough leaves suitable for matting (5). A varie- gated form of the same species (syn. P. variegatus Mia.) is always sterile and has when young longi- tudinal yellowish-banded leaves. The fatty meso- carp of several species is edible or used for fla- vouring either cooked (necessary for destroying oxalate needles) or raw. In West Malaysia it is used only by certain local tribes but in New Guinea and the Pacific (13) it represents often a substantial part of the native diet; the red fat is rich in pro- krauelianus K. ScuH. & P. terrestris WARB. belong to these fat producing species. Leaves of pandans are commonly used for thatching; the leaf-fibres vary in durability and toughness according to species (10, 11). The leaf strips or their fibres are extracted in some species and used for making sails, hats, belts, streamers, tobacco-pouches, etc. The young leaves and tip of the stem of some species are eaten. Some are also used medicinally or for magic pur- poses. The growth of pandan is mostly rapid; the wood keeps soft and juicy. Inthe Botanic Gar- 6 FLORA MALESIANA [ser. I den, Buitenzorg, P. papuanus SOLMS was planted in a pot; its stem measures now 19m height, 40 cm diam.! In pandan groves there is little under- growth and the soil below them is mostly as bare as that under bamboos, though pandan does not give as much shadow as bamboo does. A strongly Fig. 5. Planted grove of P. jiulianettii MART. mixed with an other species of similar appearance, per- haps P. macgregorii MART., cultivated for leaves and oily seeds. Nemodi, E. slope of Mt Tafa, Centr. Div., Papua, ca 2400 m alt. (ARCHBOLD) developed root system in the upper soil may be responsible. One of the reasons for its gregarious occurrence is obviously the large clumps of fruits which are always found in pandan groves. Probably ground animals such as pigs are fond of them and contribute to local dispersal. Contrary to WILLIS’s statement (12) that ‘Pandanus grows almost entirely (1) Unfortunately its age is not known. under the uniform conditions of seashores and marshes’ the genus displays an almost universal adaptability to a variety of conditions. They are known from the beach, swift streams, swamps, peat forest, mixed dry-lowland forest, limestone cliffs, savannahs, craters, mountain forests, from sea level to 3225 m. Species which grow gregariously always predominate in more or less open places and may form seres in partly devastated orcleared areas. The backward state of taxonomic knowledge of this most fascinating group of plants prevents me from enumerating predominant species by name, and therefore I have taken them together according to habitat. I have refrained from using native names, as these are local, numerous, and shifting as the sands of the sea. LITERATURE. (1) Kurz, Journ. Bot. (1867). (2) Sotms LAUBACH, Linnaea (1878). (3) WARBURG, Pfi. Reich (1900). (4) Cf. list in Webbia 4 (1913-14). (5) BACKER, Handb. Fl. Java pt 1 (1925) 42. (6) T.K.N.A.G. 52 (1935) 367. (7) De Trop. Na- tuur 26 (1937) 72. (8) BACKER, De Trop. Natuur 9 (1920) 179. (9) Wanderings (1904) 395. (10) HEYNE De Nutt. Pl. Ned. Ind. 1927. (11) BuRKILL, Dict. Ec. Prod. Mal. Pen. (1935). (12) The course of evolution (1940) 157. (13) BRown, Bern. P. Bish. Mus. Bull. 84 (1931) 30; HEDLEY, Austr. Mus. Mem. 3(1896) 1-71. (14) Journ. Arn. Arb. 20(1939) 160-186; ibid. 21 (1940) 169-175; FAGERLIND, Svensk. Bot. Tidskr. 34 (1940) 1-6. (15) MERRILL, Govt. Lab. Publ. 29 (1905) 6; Exmer, Leafi. 1 (1906) 80. 226. Littoral pandans.—Fig. 6. (A) The most common pandan of the sandy or rocky beaches is P. tectorius PARK., a rather low, and often considerably branched species (fig. 6). It is very polymorphous, was described under several specific names, and has © varieties from the Mascarenes to the Pacific. (Syn. P. fascicularis LAMK., P. odoratissimus L. f., P. samak HASSK., P. laevis KTH., P. moschatus Miq., P. variegatus Miaq.). There are a number of fixed races selected by natives, some of which were mentioned above. The plant often forms a rather narrow but dense barrier 3-7 m high, difficult to penetrate behind the pescaprae-formation of the beach proper. As such it properly belongs to the Barringtonia-beach- forest, and it is often found mixed with Scaevola, Sophora, Tournefortia, in that habitat. If dunes are formed behind the beach it extends into them. This is figured and described by BooBerG (1) from the older parts of the dunes near Poeger, SE. Java, where it is associated with Spinifex, Hyptis and Lantana. It is probably well-adapted to shifting sands. It is nearly always present on a sandy or rocky coast, and an enumeration of all localities is superfluous (5). LAM (2) mentions it from Mi- anghas island, I myself found it in narrow fringes many km long on the beaches of S. Preanger (near Oedjoeng Genteng), near Noesa Kambangan, Ban- joewangi, and inS. Bali. In P. Enggano(S. Sumatra) RAPPARD found all around the island a pandan fringe partly consisting of this species mostly 20-— 30 m, sometimes 100 m broad. Acc. to BRASs it is vol. 2] Sample treatment 7 a common strand species of the coast of the Gulf of Papua often gregarious in large numbers. It occurs also on coastal cliffs (cf. No 229). In the Pacific it seems to extend sometimes from the beach inland and to be able to colonize and form a sere on deforested areas. Fig. 6. Fruit-head of P. tectorius SOL. on the beach of SW. Java, ca X '/10. (B) P. polycephalus LAMK. is a small (2-3 m) branched species with erect spadix, capitulum con- sisting of 3-8 fruit receptacles. Known from Banka, Borneo, Celebes, Moluccas and New Guinea along sandy or rocky beaches and near non-muddy estuaries (3). According to WARBURG (4) it forms small low scrub in the Moluccas and New Guinea (5). (C) P. bidur JUNGH. (P. pacificus VEITCH) is one of the tall sparsely branched pandans (8-20 m) of beautiful stature closely allied to P. dubius SPRENG. According to BACKER (3, p. 39, 5, p. 188-189) it is found in extreme SW. Java and on the Thousand Islands (N of Batavia), on sandy or rocky places on or near the beach, sometimes locally abun- dant (5). According to BACKER (6) it also occurs in the Moluccas (Halmaheira) and W. New Guinea (Sorong). Fibre from the roots is used for thread and the leaves for thatching. The large species from P. Enggano is this one or the following. (D) P. dubius Spr. (P. latissimus Bv.). Large species averaging 12-13 m, but up to 25 m high, allied to P. bidur JUNGH. from the Moluccas and New Guinea, grows along the beach and is much valued for fibre and thatching; probably the same species as mentioned by TEYSMANN (7) as occurring by the thousands on and behind the beach in Groot Banda; also in Ceram, Ternate, &c. (E) P. labyrinthicus Kurz. Strongly branched, 3-6 m tall, many-stemmed species with numerous roots forming impenetrable tangles of stems and aerial roots (4, fig. 2) and therefore easily forming small dense groves. Known from the sandy and rocky beaches of the Westcoast of Sumatra, Sima- loer Island, West Borneo, and prob. SE. Java (3 p: 44,5; "p: 165165;p2 132): LITERATURE. (1) Hand. Se Ned. Ind. Natuur- wet. Congr. Soerabaja (1928) 377, profile and map. (2) Miangas etc. (1932) 27. (3) BACKER, Handb. Fl. Java pt 1 (1925) 45. (4) Pfil. Reich (1900) 20. (5) BAcKER, De Trop. Natuur 9 (1920) 178-191; 10 (1921) 12-17. (6) in HEYNE, De Nutt. Pl. (1927) 129. (7) Natuurk. Tijdschr. Ned. Ind. 23 (1861) 356. 227. Littoral mudbank pandans. Only one record is known to me of a gregarious species peculiar to tidal mudbanks of the Fly River up to Ellengowan extending to 320 km from the sea (1). LITERATURE. (1) BrAss, Journ. Arn. Arb. 19 (1938) 178. 228. Coastal forest pandans. No species except scattered specimens of the beach species have been recorded from the coastal forests. 229. Coastal cliff pandans. It appears that the coastal cliff pandans have no preference for a special type of rock; they occur both on elevated coral reefs and other limestone, and on andesite. (A) P. tectorius PARK. Creeps up from the beach onto the rocks. Mentioned e.g. by CoerT from Popoh, S.Central Java, (1), Kroe, SW. Sumatra. (B) Pandanus spp. On a limestone terrace be- tween Waren and Wariap, Miss Grpps passed through a striking group of old Pandanus 30m tall; Papuans said this was also in the forest in isolated groups. Miss GiBBs wrongly supposes them to represent the original vegetation on the limestone later displaced by more rapidly growing mixed forest trees (2). LANE PooLe figures a beauti- ful stand of Pandanus on coastal limestone (3). LITERATURE. (1) De Trop. Natuur 22 (1933) 80. (2) Flora & Phyt. Arfak Mts (1917) 18. (3) Rep. For. Res. Terr. Pap. Nw G. (1925) fig. 22. 230. Lowland open swamp pandans. There is every transition between wooded swamps (swamp forest) (No 231) and open swamps, but for the sake of convenience the open swamps are described separately. Riverine pandans(No 235) may sometimes fill cut windings of rivers and resemble swamp pandans. Few gregarious swamp pandans have been named. (A) P. radula Wars. Described from Sumatra, but recorded and figured by Hus. WINKLER from the Bornean ‘danaus’ (1). (B) Pandanus sp. div. BEccARI records from Bor- neo some pandans which form impenetrable thick- ets in marshy localities (2). In South Sumatra, DE Voocp found in the rawahs of Sekanak, near Palem- 8 FLORA MALESIANA bang, a creek connection with the river Musi filled with pandans(3). InE. CelebesI found, S of Palopo, Bone, marshes locally dominated by a rather tall pandan with beautiful red fruits (STEENIS 10367) locally in rather pure groves in old lowland second- ary forest. In the Fly river area BRASS and RAND mention from Gaima pandans in dense stands on mud banks (4). LITERATURE. (1) Das Leben der Pflanze 6 (1913) Abb. 79, p. 232. (2) Wanderings (1904) 395. (3) Trop. Natuur 21 (1932) 64. (4) Bull. Amer. Mus. Nat. Hist. 77 (1940) 361. 231. Lowland swamp and peat forest pandans. Of this group also few species are sufficiently known. In a swamp forest no real peat layer is pre- sent; when peat forest is mentioned layers of more than 1 m peat are present. (A) P. setistylus Wars. According to WARBURG (1) a lowland swamp species in NW. Guinea. (B) P. papuanus SoLms LAUBACH. Large species, stems 10-15 m, prop roots developing below the branches; all along the E.coast of New Guinea and adjacent islands; gregarious in locally extensive groves on permanently marshy soils along the coast and along rivers and lakes. Papuans use the fibre from the 7-8 m long prop roots. Cultivated in Halmaheira where the leaves are used for mat- ting; not useful for plaiting (2). (C) P. atropurpureus MERR. & PERRY. Large species 14-16 m tall, with branched crown and long stilt roots; fruit head 43 by 20cm; collected by Brass, ‘plentiful in the more open and swampy parts of floodplain rain-forests’ in the Idenburg river region, New Guinea; also collected at 850m occasionally in rain forest on slopes (3). (D) P. johorensis Mart. Shrub 4m tall, from the Mal. Peninsula and East Sumatra, in the low- land swampy forest, gregarious (2, p. 132). (E) P. exiguus MERR. & PERRY. A dwarf species, not branched, found by BRAss gregarious in con- siderable numbers in forest undergrowth round the edges of a swampy depression and extending to the surrounding low ridges, Palmer river, Papua (9). (F) Pandanus spp. ‘Bengkuang’ is in E. Sumatra, Bengkalis Isl. and the opposite inland of Sumatra typical of the acid soils and central parts of the peat forests; together with ‘linau redang’ (prob. the palm Cyrtostachys renda BL.), it is dominant over a vegetation of sedges Scleria &c. in a ‘fairy forest’, according to J. H. p—E HAAN, Febr. 1935. ENDERT (8) found the same in the Palembang peat forest where a very large pandan is associated with Eugenia, Tristania and Architaea. In Borneo, BEc- CARI (4) found along the Sumundjang river a ‘terri- ble Scleria’ covering large tracts of swampy land with impenetrable thickets in which a Pandanus is a conspicuous feature; he found also flooded forests with masses of the sedge Mapania and Pandanus as a substage (4, p. 349). In Obi Island, Moluccas, G. A. L. DE HAAN took a photograph of a swamp forest at 200m on serpentine rook, E of lake Telaga, in which Pandanus was abundant (5). In New Guinea, LANE PooLe records from the Ramu valley two species of Pandanus associated with fserz I Sarcocephalus and Saccharum spontaneum L. This is the community of the true swamps, depth a few in. to 4 ft, underneath mud (6). BRAss & RAND recorded from Sturt Isl. in the Fly river a low species forming a substage layer over considerable areas which are frequently flooded (7). LITERATURE. (1) Pfl. Reich (1900) 81. (2) Herne, De Nutt. Pl. Ned. Ind. (1927) 132. (3) Journ. Arn. Arbor. 21 (1940) 173. (4) Wan- derings (1904) 348-349, 395. (5) Tectona 31 (1938) 170, fig. 2. (6) Rep. For. Res. Terr. Pap. Nw G. (1925) 59. (7) Bull. Amer. Mus. Nat. Hist. 77 (1940) 361. (8) Tectona 13 (1920). 125. (9) Journ. Arn. Arb. 20 (1939) 171. 232. Inland rock and cliff pandans. Only records of non-identified species are on my list. On dry tuff-ridges near Sibolangit and on the margin of the limestone sinter terraces of the hot springs Pandanus is rather gregarious and con- spicuous (1); it is clearly more tolerant than other species of the surrounding forest. F. C. VAN HEURN mentioned the conspicuous occurrence of pandans on the limestone sinter terraces of Tinggi Radja, Sum. Eastcoast (4). WiTKAmp found in the inland of Koetei, E. Borneo, P. on limestone cliffs (2). WALLACE mentioned in the Key Isl. pandans and Dracaena characterizing the vegetation of the more rocky places (3). Brass found P. hollrungii WARB. in Papua, Palmer river, ‘apparently restricted to poorly drained soil on flat ridgetops, ca 100 m alt.’; small species 4—5 m tall (5). For rocks and ridges at high altitude see No 266-268. LITERATURE. (1) Natuur in Indié (1937) 13. (2) T.K.N.A.G. 46 (1929) 209. (3) Mal. Archip. ed. II, 2 (1869) 111. (4) Studién etc. (1923) 77-83. (5) Journ. Arn. Arb. 20 (1939) 160. 233. Pandan in savannahs. Pandans are sometimes tolerant of burning prac- tices and resistant to some extent to deforestation. For these reasons pandans are sometimes a striking feature in savannahs. P. ovatus KuRz is a dwarf species growing in tufts in open grassy places and on hill-tops in S.Siam and the Mal. Peninsula as far S as Central Johore (6). In Celebes the SARA- sINs (1) found S of the Topapu Mts burned grass- land with stretches of burned young forest; groups of a candelabra-pandan had survived in the charred area. ARCHBOLD & RAND record (2) from Dogwa, 40 m alt., 99 km from Daru, Papua, flat valleys to the W of Dogwa, covered with tall dense grass and scattered low trees, or with extensive stands of pandans. The ridges were also grass- covered, some dotted with low trees, some with stands of Pandanus. The clear stands of 6-10 m high might be called ‘pandan-forest’ and were a striking feature. The stands extended over some of the valleys in which the ground was very wet and also onto the ridges where the soil was dry and where burning of the grass was practised by the natives. W of Merauke, S. Dutch New Guinea, a similar open savannah was found by H. NEvER- MANN (3) showing a sparse association of Pandanus and Melaleuca. J. W. R. Kocu figures the same vol. 2] Sample treatment 9 from the neighbourhood of Merauke (4); a species with thin tall stems, candelabra-branches and hang- ing foliage. Father H. GEURTJENS mentions it from about the same area where he found an anthro- pogenic savannah with termite (‘ant’) hills, scrub and pandans in extensive flats behind the beach and dunes on hard grey loamy soil (5). In Daru Isl., W. Div., BRAss collected a variety of P. tecto- rius PARK. aS abundant in the substage and con- spicuous in the savannah forests as a tree 7-8 m high (7). He found the same species in the Wassi Kussa area in poorly drained savannah forests: mature trees were rare, but seedling plants were very abundant in places (7, p. 164). P. brassii MART. is recorded by Brass from Dogwa, Oriomo river, West. Div. Papua, as commonly scattered over open grass-slopes, and also forming patches some acres in extent of almost pure forest of trees 4-5 m tall on seasonally wet hummocky ground in savan- nahs (7, p. 165). LITERATURE. (1) Reisen in Celebes 2 (1905) 119. (2) Bull. Amer. Mus. Nat. Hist. 68 (1934) 578. (3) Bei Sumpfmenschen und Kopfjagern, 2. Aufl. prob. 1936, p. 23, 37, fig. p. 48. (4) ZW.N. Guinea Exped. 1904-05 (1908) 503, fig. 106. (5) Op zoek naar oermenschen. Roermond (prob. 1934) 28-30, 53, fig. (6) BURKILL, Dict. Ec. Prod. Mal. Pen. (1935) 1650. (7) Journ. Arn. Arb. 20 (1939) 163. 234. Dry lowland forest pandans. Except in streams, swamps, on ridges and rocks, near solfatara and hot springs, I have no data on gregarious occurrence of pandans; scattered speci- mens and small colonies of pandan are found in nearly every type of lowland forest on dry soil, specially as substage plants. P. kaernbachii WARB. was found in Sturt Isl., Lower Fly, abundant in the substage of the flood-plain rain forests by BRASS (stem 8-10 m long) (1). LITERATURE. (1) Journ. Arn. Arbor. 20 (1939) 161. 235. Riverine pandans (pandan rasau or resau).— Fig. 7. Along rivers and swift streams, both freshwater and peatwater, all over the Archipelago special species of Pandanus may form a conspicuous fringe on the banks. The stems are mostly submerged. Sometimes rivers are overgrown by them and the passage is obstructed; and isolated river arms are rapidly filled with pandan, specially in the Malay Peninsula, Sumatra and Borneo. This is not known from Java! In the Malay Peninsula RIDLEY (1) mentions P. helicopus KURZ, a slender species up to 6 m tall, forming dense thickets often for miles and blocking the waterway in rivers. P. prainii MART., a small species, stem 2!/2—-31/2 m tall, is found in small water holes with the top of the fruit just above the water. P. immersus RIDL. with erect leaves forms equally dense thickets in streams, P. militaris R1DL. is another riverine species. In Sumatra TEYSMANN (2) found a gregarious riverine pandan near Sibolga, along the river which debouches near Djagadjaga in Tapanoeli Bay. In the Koeboe regions, in the Lalang river system, N of Palembang P. helicopus Kurz, up to 6 m tall, mostly only 2 m emerging from the water, known from the Mal. Pen., Sumatra and Banka, com- monly forms a fringe 5-10m broad along the rivers (3). Acc. to ENDERT(12) the peat water rivers of Palembang are fringed by three different species Fig. 7. Riverine rasau pandan in fruit, NE. Borneo. (ENDERT) of Pandanus, amongst them P. helicopus. In the less swiftly running water a floating fringe of Susum is found in front of the pandan zone. Miss E. POLAK (13) also photographed them inSE. Sumatra (fig. 7). In Banka TEYSMANN found a riverine speci- es in the Djeboes flowing into the Kampa estuary, and one in the river Kepo above the brackish water level; natives sometimes cut the vegetation to keep the water open (4). BECCARI peculiarly does not mention the striking riverine pandans which occur in Borneo. v.D. ZWAAN found them in SW. Borneo, in the Sg. Rasau Lemandau, Kota Waringin (5). In E. Borneo ENpertT figured them along the Kom- beng stream from the Upper Mahakam system (6) where he found two species, and H. WITKAMP recorded them from the Sg. Koetai-lama there. From SE. Borneo Hus. WINKLER (14) mentions them. From New Guinea there are several records. HELDRING (7) found them in the peatwater of the De Wildeman river, upwards of ca 100 km from its confluence with the Eilanden river; the depth of the thick pandan fringe exceeds 6 m. Also along the Lower Digoel pandan swamps are present. H. WITKAmpP found them in some cut off bends of the 10 FLORA MALESIANA [ser. I, vol. 2] Digoel above Tanah-merah (1939). The same was observed by SALVERDA (8); he photographed rows of a large pandan on the bank of the Bloemen river. Brass & RAND (9) found on the river flats a gre- garious pandan at 100 m alt., at Palmer Junction Camp, Central Fly, SE. New Guinea. LAM also mentioned riverine pandans from NW. New Gui- nea (10) and LorENTz gave a good figure (11). Brass found P. pseudopapuanus Mart. as a tall tree up to 25 m or more near Wuroi, Oriomo river, W. Div., plentiful in riverbank rain-forest (15). P. leiophyllus MART. was found by Brass in the Middle Fly in rain forests; in the Lower Fly region he found it gregarious in extensive, dense, pure stands, 6-8 m high, as undergrowth and substage in tall forest of Bruguiera sexangula (Lour.) Por. and Mpristica hollrungii WARB., on mud flats frequently inundated by fresh water backed up by the tides, on Sturt Isl. and vicinity, Papua. In the Upper Wassi Kussa river it was restricted to riverbanks in the rain forests (15, p. 171). P. aggregatus MERR. & Perry forms clumps 12-14 m tall, of several trees rising from a small compact group of erect prop roots, stems branched or not; abundant in the substage layer of riverine forests at 100 m alt. (15, p. 176). A common species in Papua is P. lauterbachii K.ScH. & WaARB., found by BRAss gregarious in extensive communities forming un- dergrowth of low substage 4-14 m high, in tall forest of river flood banks, Lower Fly about Sturt island, Upper Fly on river lowlands, and near Bern- hard camp, Idenburg river (in Jitt.). LITERATURE. (1) Flora Mal. Pen. 5 (1925). (2) Natuurk. Tijdschr. Ned. Ind. 14 (1857) 358. (3) Heyng, De Nutt. Pl. Ned. Ind. (1927) 131. (4) Natuurk. Tijdschr. Ned. Ind. 32 (1873) 40, 77. (5) Report 1938. (6) Midd. Oost Born. Exp. (1927) 220, 221, fig. 80, 257, 239, fig. 90. (7) Jaarb. Mijnw. Ned. O. I. Jg. 40 (1911) Verh. p. 78-79. (8) Rapp. Orient. Expl. ZW. N. G. 1936-7, mimeogr. p. 19, phot. 31, p. 24. (9) Bull. Amer. Mus. Nat. Hist. 77 (1940) 368. (10) Fragm. Pap. IV, 205-208. (11) Zwarte menschen, witte bergen (1913) p. 121 photo. (12) Tect. 13 (1920). 125. (13) Proc. Akad. Wet. A’dam 30 (1933) no 3. (14) Bot. Jahrb. 50 (1914) 196. (15) Journ. Arn. Arb. 20 (1939) 165. 236. Pandans in mountain forests on volcanoes, and near craters.—Fig. 1-3, 8, 10. The wide edaphic tolerance of pandans which enables them to grow in acid peat swamps, on brackish seashores, and on limestone cliffs allows some species also to grow gregariously on vol- canic slopes and summits and near craters, where they often form a conspicuous feature. The same species are found in all these habitats but they are more gregarious near the craters. VoLz (1) found pandan a characteristic plant in the Batak Lands, W.Central Sumatra between 1600 and 2000 m. He also mentions it (same species?) to be abundantly planted for plaiting (/.c. 82) in the Karo Lands. On Mt Sinaboeng he found towards the summit (J/.c. p. 5-6) low scrub of Rhododendron and pandans. According to BACKER this is P. platycarpus BACKER ms. (non WARB.) which is also found gregariously on Mt Sibajak and near Wilhelmina waterfalls. MARTELLI mentions P. vriensii MART. from Mt Sibajak. BARTLETT (2) saw a belt of P. sumatranus Mart. on Mt Sinaboeng between the forest belt and the ericoid scrub, between 1700-2300 m (vern. toemenang) (7). KEMMERLING (3) says that the ee Fig. 10. P. furcatus Roxs. in a mountain swamp, above Buitenzorg (Telaga Saat), ca 1400 m alt., ca X ‘1/15. gradually increasing frequency of pandan indicates the approach to the forest limit on Mt Talamau (= Mt Ophir) (6). On Sorok Merapi pandan is very frequent between 2000 and 2145 m (summit); trees are often covered by volcanic ash. On Mt Tandikat he found (/.c. p. 19, 27) ferns and pandan in ash and sand of the 1889-eruption. From the Goudberg, E. Atjeh, VoLz (4) mentions abundant pandans round the solfataras of Van Heutsz crater at 650 m alt. LEKKERKERKER (5) mentions the same; Pinus merkusii is also found there. On Mt Dempo, Bencoolen, Forses (8) found at 2100 m patches of tall pandans on the sides of a gorge but nowhere else on the mountain. Mt Kaba, also in Bencoolen, a much lower active volcano, however, has a crater very rich in pandans, he says (/.c., p. 228) above 2250 m associated with Melastoma and Lonicera; these are figured by DE Voocp (9), (fig. 8), forming impenetrable thickets; elephants feed on the young tufts of foliage and the fruit. In Java gregarious pandans are rare in the mountains: a species occurs gregariously in Central Java, Kedoe, on the Ran- djak near the summit of Mt Andong dcc. to ALTONA (10). On Mt Pandan, Madioen, pandans are frequent in undergrowth, 700-900 m. ZOLLINGER found a large species gregarious on Mt Taroeb, the older part of Mt Lamongan, E. Java (11). This is Sample treatment of vol. 3 MALAYSIAN PLANT GEOGRAPHY BEING A FLORISTIC AND GENETIC PLANT GEOGRAPHY OF THE MALAYSIAN REGION ILLUSTRATED BY NUMEROUS MAPS BY C. G. G. J. VAN STEENIS Contents: Ist PART: Floristic plant geography Chapter 1. Introduction to the aims and methods of floristic plant geography. Chapter 2. Historic review of the plant geographical theories advanced for the Malaysian region or parts thereof. Chapter 3. Tabulated analysis of the distribution of the genera of Malaysian Phanerogams. Chapter 4. List of the genera recorded from Malaysia, with their synonymy and distribution. Chapter 5. Analysis of the floristic standing of the islands and island groups separately. Chapter 6. Attempt towards floristic synthesis and proposals of a new division of Malaysia into provinces and districts based on the distribution of Phane- rogamic genera. N.B. The preparation of this part of volume 3 is in an advanced stage. The second part which will deal with the genetic plant geography is in the initial stage. The complete volume will comprise appr. 600 printed pages.—ED. FLORA MALESIANA [ser. I THE DELIMITATION OF MALAYSIA AND ITS MAIN PLANT GEOGRAPHIC DIVISIONS as based on the distribution of the genera of Phanerogams Plant geographical demarcations are sharp for one single genus, one species, &c. but the general pic- ture is far from clear. This is quite natural as the present vegetation represents only the outcome of continuous plant-migration and subsequent floras in past aeons. Everywhere progressive elements thrive together with relics. The localities of the lat- ter are but rarely connected by phyto-palaeontolo- gical localities proving their former occurrence and migration and, possibly, elucidating their present disjunct distribution. An example from the Malaysian lowland is Dryo- balanops, a camphor producing genus of Diptero- carpaceae consisting of 7 species and occupying a coherent area in Central Sumatra, the Malay Pen- insula and Borneo ( Fl. Mal. I, 4, p. ly, fig. 44). In the Late Tertiary Period it thrived in the ad- joining parts of S. Sumatra and W. Java, where now large quantities of silicified timber of unques- tioned identity testify of its formerly frequent oc- currence where it is at present entirely extinct. In N. Sumatra I found on the high mountains com- munities of the sedge Schoenoxiphiumkobresioideum KUxk. with great numbers of Ericaceae, the latter so well represented numerically that they strongly suggest a recent development. Schoenoxiphium is, however, according to Dr KUKENTHAL, the ‘mis- sing link’ between the African Schoenoxiphium and the Asiatic Kobresia; the N. Sumatran species is the first of its genus to be found out- side Africa. The segregation of the present mixed Malaysian flora could be accomplished by determining in each genus its centre(s) of development in order to evalu- ate the present distribution in Malaysia in respect to the whole generic area. In combining the figures of the several genera, the laws governing the past and present distribution (genetic or historical plant geographic method) of the Malaysian flora might be traced. Small groups are particularly suited to this method. Another way to analyse and demarcate the Ma- laysian flora is the floristic or quantitative method by which the single groups are not valued individually. Its sole purpose is to trace demarcation lines where the botanical ‘melting-pot’ shows a more or less abrupt change, or, a threshold in its com- position. In following the latter method here J have adopt- ed the genus as the unit for distribution or, in some cases, a well-marked subgenus or section. To study the distribution of the species would be impossible, their known synonymy and acknowledged area being generally far less reliable than those of the genera. Similar attempts have been made formerly for local areas based on special collections, e.g. by WarburG for the Papuan flora (1891), by STAPF for the Kinabalu flora (1894), and by MERRILL for the Philippine flora (1923). Biological lines have been drawn in and around Malaysia by the dozens based on zoological, pa- laeontological, or botanical arguments. Similar lines have been derived from palaeogeographical, clima- tological, and geological sources or combinations of these. The lines are partly of a connecting, partly of a dividing character. Several efforts have been made both by zoologists and botanists to co-ordi- nate the lines. Striving towards a synthesis, bota- nists often derived their conclusions from a small or at any rate limited group of related plants and tried to make their results agree with one of the more than two dozens of theories proposed for Ma- laysian geology. HALLIER’s work is both the most daring and the least sustained by facts, MERRILL’S the best provided with basic material. Is is striking that though MERRILL’s botanical results distinctly show the overwhelming West Malaysian character of the Philippine flora, he prefers to put his results into the frame of a geologic division of the Malay- sian Archipelago in two continental shelf-areas separated by an unstable region. ; As the various natural sciences mentioned above show different aims and means, it seemed to me that the first object of Malaysian plant geography must be to sift and analyse the botanical facts them- selves and to synthesize the major botanical fea- tures, without binding them to some geological or otherwise non-botanical theory. Malaysian botany has its own problems. An example is the different degree of alliance between the floras of different altitude: the mountain flora of Java is nearly identical with that of most of Su- matra but very different from the Bornean moun- tain flora. The lowland floras of Sumatra and Borneo are practically identical, however, but widely differ- ent from the present Javan flora. The lowland flora of the Philippines is closely allied to that of Borneo, but the upland flora of N. Luzon shows a remark- able set of E. Asiatic genera and species. The low- land flora of New Guinea is essentially Malaysian but the summit flora has produced a unique Austra- lian-Subantarctic element. The present study is based on several years of research in the Buitenzorg Herbarium, on litera- ture, and on accumulated herbarium material. Many unpublished data have been used, especially those from the numerous large collections made in the years 1930-1940. I am of opinion that, though unexpected, the outcome is both clear and well founded. Some predictions founded on my results have already proved to be correct, which gives them marked support. The results are based on figures derived from more than 2100 distribution maps of all genera of Phanerogams native in Malaysia. For each genus I have tried to locate the centre(s) of specific development. This has led to an ar- rangement of these genera in 5 types which gave the following figures: vol. 3] Sample treatment 3 F ne Sos 30 Se) Ore Number Percentage Different types of generic distribution: 7 of eenera | GE totale Tyre 1. Occurring in Asia, Australia, and Malaysia; no distinct centre in the paleo-tropics 602 Ds Tyre 2. Centre of specific distribution clearly i in ee absent or scarcely represented in Australia 574 26.3 iiveEnes. Centre ol development i in Malaysia, and some 2 outposts ' in surrounding regions 380 ) 26.6 | Type 3a. Genera known only from one island or island group in 876 40.4 Malaysia (endemic genera) 296 ) 13.8 ( Tyre 4. Centre of development i in Australia: absent or r scarcely represented in Asia 94 4.3 Type 5. Centre of development in the Pacific- Subantarctic region 32 1.4 Total | 2178 100.1 Though the census was made in 1945 and, there- fore, the ultimate figures will be slightly different, they are so well pronounced that their essential tendency may be seen as final. A first characteristic is the high percentage of type 1 which could be termed the ‘Indo-Australian basic flora stock’. Another remarkable feature is the great number of Asiatic genera absent from or only just reaching the Australian continent, which shows that the old opinion of HOOKER & THOMSON, MIQUEL, and ZoL- LINGER of the close alliance between the Malaysian and the Asiatic floras holds. The high percentage of genera entirely confined to Malaysia (type 3a) or centering in Malaysia (type 3) together comprising 40°o of the total proves that the Malaysian region is worthy of the rank of a separate plant geographical province on an equal footing with that of SE. Asia. The endemic genera are far from being equally distributed over the island groups, and certain is- lands are much richer than others. The grouping of the endemic genera is shown on a map (fig. 1). The Australian resp. Pacific elements play, meas- ured by figures, a very unimportant role in the general picture. Their number was formerly much overestimated owing to the occurrence of some con- spicuous plants such as Araucaria, Eucalyptus, Ca- suarina, Banksia, Grevillea, and Acacia. This was specially suggested for the New Guinean flora, and has even made a geologist declare that the New ‘Guinean flora possessed essentially an Australian character. Our knowledge of the Papuan flora is still incomplete but there is no reason to assume that it is more incomplete as regards the Australian element than as regards the Asiatic-Malaysian ele- ment. A most important point is the delimitation of the wegion we have defined as the Malaysian region, ‘and:accepted as a natural plant geographic unit. It has 4.contacts or ‘bridges’ with the adjacent floral regions, viz with Asia in the Malay Peninsula and in ‘the Philippine Islands, with the Pacific islands in the Bismarcks and Solomons, and with Australia in the island of New Guinea. Is there any abrupt demarcation in the generic composition or is the change in the flora gradual? 1. The Malay Peninsula. Though the NW. fron- tier of Malaysia is situated on the isthmian land- connection with continental Asia in the Indo-Chi- nese Peninsula, the Malay Peninsula has up to the north, approximately near the line Alor Star—Sing- gora (that is a little north of the political border), a typical Malayan flora intimately allied to the floras of Sumatra and Borneo but differing strongly from that of Indo-China. On the other hand, the immediate neighbourhood of the latter has not appreciably enriched the Malay Peninsular flora with continental elements if compared with the Fig. 1. Number of endemic genera of Phanero- gams in the several islands and island groups of Malaysia, according to a census made in 1945. Sunda Islands. The figures are: the Malay Penin- sula has 36 Asiatic genera not found elsewhere in Malaysia, Sumatra /8, Java 27, and Borneo /0. Conversely, the Malay Peninsula shows 196 Ma- laysian genera which are absent from continental Asia, or have only a stray record in Ceylon, Pegu or Siam. The close connection between the floras of Sumatra and Borneo when matched with the flora of the Malay Peninsula is illustrated by the 4 FLORA MALESIANA [ser. I fact that the islands together possess only 62 Ma- laysian genera which have as yet not been found in the Malay Peninsula. To the ‘generic pressure’ from the Malaysian side towards Asia must be added the endemic genera of Malaysian affinity found in Su- matra (17), the Peninsula (4/), and Borneo (59), that is //7 in all. The total of Malaysian genera on the Malaysian side of the NW. frontier becomes thus 196 + 62 + 117 = 375 genera. On the continental side of the frontier are about 200 Asiatic genera, recorded from Burma, Siam or Indo-China which have not been found in Malaysia. The ‘demarcation-knot’? Malaysia <— SE. Asia amounts therefore to 375 + 200 = ca 575. In oth- er words: 575 genera respect the demarcation Alor Star—Singgora.—Cf. fig. 2. I employed the same method for the other Ma- laysian frontiers. The magnitude of the ‘demar- cation-knots’ is a means of measuring quantitati- vely the borders and ‘knots’, both of the Malaysian frontiers and the divisions inside the Malaysian flora. Knots of equal magnitude delimit districts or provinces of equal standing, and indicate the stand- ing of the enclosed area. 2. Philippines. An analysis of the Philippine flora according to this method shows the occurrence of 32 genera of type 2 (Asiatic genera) in the Philip- pines (among these 23 are found only in Luzon and 21 genera are mountain plants!) which are not re- corded from other parts of Malaysia. This is about of the same order as the figures found for Borneo, the Malay Peninsula, and Sumatra. It confirms the essentially Malaysian character of the Philippine Fig. 2. The 3 principal floristic ‘demarcation knots’ of the Malaysian flora. flora. In the Philippines occur 388 Malaysian gene- ra which have never been found in Formosa or the adjacent parts of China. The Philippines possess 33 endemic genera. Of the 1185 genera known from Formosa, 265 are not found in the Philippines. The ‘demarcation-knot’ Philippines<—>Asia is thus 388 + 33 -+ 265 = 686. This is slightly larger than in the Malay Peninsula, but still of approximately the same rank.—Cf. fig. 2. 3. New Guinea. For the island of New Guinea compared with Queensland the figures are as fol- lows. In New Guinea 175 Asiatic genera have their most eastern distribution; further the areas of 345 Malaysian genera! end here and, in addition, there are 124 endemic genera which is a total of 644 Fig. 3. Numbers of eastern-centred genera in the Archipelago; above the hyphen: Australian genera, below it: Pacific-Subantarctic genera. genera. In Queensland and North Australia about 340 Australian genera, absent in New Guinea or other parts of Malaysia, occur. The total of the ‘demarcation-knot’ in Torres Straits is thus 984. This is markedly larger than the figures pertaining to the Malay Peninsula and the Philippines. Here is, apparently, one of the main demarcations of the Palaeotropic plant world.—Cf. fig. 2. The analysis of eastern genera, Australian, Pa- cific or Subantarctic (types 4 and 5) in Malaysia shows that New Guinea possesses 40 of those not recorded elsewhere in the Archipelago. Of these, 16 are confined to savannahs, 13 to the forest, 10 to the high mountain summits, and 1 to the man- grove swamps. New Guinea has certainly the largest number of eastern genera in Malaysia but not many more than elsewhere in the Archipelago, as may be concluded from the map reproduced in fig. 3, which shows a rather gradual decrease of eastern genera towards the west. It seems natural that New Guinea with its enormous and varied land surface and geograph- ic situation should harbour most of them. 4. Bismarcks, Solomons, and New Hebrides. Here I cannot find figures of the magnitude of the other ‘knots’. This is partly due to the backward state of knowledge of these floras (both as regards col- lecting and literature), but the main cause is that the Micronesian and Melanesian floras are poor, (1) Of some genera which are, in New Guinea, richly developed, e.g. Saurauia, Medinilla, Rhodo- dendron, Polyalthia, Cyrtandra, etc. with dozens of species, a stray species sometimes occurs in N. Queensland. vol. 3] Sample treatment 5 and without a distinct character in any degree com- parable with that of the floras of Asia and Austra- lia. They are mainly derivatives from the Malay- sian flora. The demarcation of Flora Malesiana against these island groups is artificial, and we know it. For practical reasons we have refrained from in- cluding these floras in the present one. These statistics show that Malaysia, as we accept it, isa natural unit, well-demarcated at its frontiers except towards Micronesia and Melanesia. The In- do-Asiatic flora rather abruptly ends in New Gui- nea. On account of a few facts but with a remarka- bly clear insight, ZOLLINGER had, already in 1857, come to the same conclusion. Inner Divisions of the Malaysian flora. In 1845 EARLE propounded his ‘bank’ or ‘shelf’ theory; he distinguished the western shallow sea covering the Asiatic continental shelf, or Sunda shelf, from a similar shallow submerged extension of the Australian and New Guinean land, or Sahul shelf; these were separated by a non-continental central part. A year later SAL. MULLER, apparently quite inde- pendently, proposed on zoological arguments a division of the Archipelago into two parts through the Makassar Straits. In 1857 ZOLLINGER extended this line northwards between Celebes and Mindanao; towards the south he did not extend the line somewhere between the Lesser Sunda Islands, but east of them. He was fol- lowed in this by MIQUEL, in 1859. In 1863 WALLACE proposed the boundary called ‘Line of Wallace’ by HUxLEy, which line has been so amply discussed up till the present time. WAL- LACE seems to have founded this line independently of ZOLLINGER; nowhere did I find any mention | made of the Swiss’s work. _ NverMEIER,! in a historical review of Wallace’s Line, has shown that WALLACE himself changed his views on the subject, and finally did not take it too seriously. Much later, MERRILL & DICKERSON projected a northern continuation of WALLACE’s line along the W. side of the Philippines, between Palawan and Mindoro, and recently v. MALM shifted WALLACE’S Line in its southern extremity between Bali and Lombok towards the east between Flores and Ti- mor. The different courses of the Line of WALLACE are indicated in the map reproduced in fig. 4. In using the method of the ‘generic demarcation- knots’ we may first calculate the importance of Sunda Straits and Java Sea between Java and Su- matra-Borneo. In 1933, I tried? to draw a compar- ison between these floras. The flora of Java ap- peared to be very poor in relation to that of the other islands: 3 (small) families and 111 genera— several of which show an abundant specific devel- opment in Sumatra and Borneo—are absent from (1) Tijdschr. Kon. Ned. Aardr. Gen. 14 (1897) 758-769. (2) Bull. Jard. Bot. Btzg. III, 13 (1933) 23-28. Java. On the other hand Java possesses many gene- ra which are absent from Sumatra and Borneo. The climate cannot be held responsible for this differ- ence as West Java does not differ in climate from South Sumatra. The genera occurring in Java but not in Sumatra and Borneo, however, belong most- Fig. 4. Various courses of the dividing line between East and West Malaysia: all have Makassar Straits in common, that of MERRILL & DICKERSON is a deviation only in the north, those of SAL. MULLER (the oldest line) and v. MALM are deviations in the S. end only. The line accepted here is that of ZOLLINGER. ly to plants bound to a two-seasoned climate. Most of them belong to type 1 or 2; they occur in the monsoon forests of Burma, Siam, and Indochina, are absent from the wet central part of West Ma- laysia, but reappear in Java, the Philippines, Ce- lebes, and the Lesser Sunda Islands. This disjunc- tion is doubtless climatic. It remains a telling fact that the monsoon plants are even in Timor mainly Asiatic, without any appreciable or significant ad- mixture of Australian drought plants. In any case, the generic ‘demarcation-knot’ of Sunda Straits and Java Sea is at least 200, which agrees with the importance the zoologist VAN KAMPEN attached to Sunda Straits. I have not been able to trace a knot of this rank between the other parts of West Malaysia: Sumatra, Mal. Peninsula, Borneo, and the Philip- pines. There is, between Java and the Lesser Sunda Is- lands, very little basic difference: the flora of the latter is characteristically a depauperized Javan flo- ra without appreciable admixture of Australian elements. I had formerly found this for the moun- tain flora.? A characteristic of both Java and the Lesser Sunda Islands is the low percentage of en- demic genera (cf. fig. 1). Java and the Lesser Sunda Islands form together a separate province of Malaysia, a view already held a century ago by ZOLLINGER and MIQUEL. (3) Bull. Jard. Bot. Btzg III, 14 (1936) 56-72. 6 FLORA MALESIANA [ser. I Turning to other parts of Malaysia, Makassar Straits proves still to be a very important line of demarcation. No less than 297 genera of Phanero- gams occur in Borneo but not east of Makassar Straits. The number of eastern genera occurring in Celebes but not in Borneo is much less well mark- ed (cf. fig. 3), the total number of eastern genera Fig. 5. Numbers of Asiatic-centred genera which are known to occur in Malaysia only in 1 island (figure above the hyphen) or 2 islands (figure below the hyphen), illustrating intensity of ‘Asiatic in- fluence’ in the Malaysian flora. being much smaller than that of western genera. For the northern extension of the Makassar Straits demarcation line there is a choice between the line separating Celebes from Mindanao (ZoL- LINGER, WALLACE) or the ‘corrected line’ between Palawan and Mindoro (MERRILL & DICKERSON). In the first place, the large number of Asiatic ge- nera which occur in only one or two islands in W. Malaysia indicates a remarkably sharp demarca- tion, and is to some degree a good measure for ‘Asiatic influence’; it clearly supports the original course of Wallace’s Line. This is demonstrated on the map reproduced in fig. 5. Secondly, on comparing the number of western and eastern elements according to the scheme re- produced in fig. 6, where A+ A’ is opposed to B+ B’, it appears that 110+ 71 = 181 genera find their eastern border E of the Philippines, but only 9+ 34 = 43 find their western border W of the Philippines. This is further evidence that the original Line of Wallace is substantially more im- portant than the corrected line. Thirdly, it must be considered that typical West Malaysian genera and families show a large spe- cific development in the Philippine Islands closely connected with the development in Borneo, as shown by MERRILL in 1923 in his valuable study on the disuribution of Dipterocarpaceae. From these arguments it can only be concluded that the ‘demarcation-knot’ in the N. part of Ma- laysia agrees with the original Line of Wallace. There are no botanical but only geological argu- ments for keeping the Philippines apart from West Malaysia. Having thus established by the quantitative method of ‘demarcation-knots’ the borders of the Malaysian flora against the adjacent areas of the Indian and Australian floras, and the main divi- sions within its boundaries (a West Malaysian, East Malaysian, and South Malaysian province (fig. 1, p. xi), it remained to divide the provinces into districts. The tentative result is reproduced in fig. 7 where the thickness of the lines indicates their phytogeographical value. The interpretation of the floristic divisions in- volves much discussion and explanation: 1°. When geological maps are examined there is a discrepancy between botanical delimitation and that of the shelves, viz in Torres Straits, Sunda Straits and the Java Sea, which are botanically very important demarcations. The Philippines— though situated outside the continental shelf—have a West Malaysian flora. The great importance of Torres Straits was pointed out already by WARBURG (1891). Accept- ing geological arguments as decisive, LAM, in 1934 and 1935, has advanced an explanation on the basis of WEGENER’s theory, and made the bold specula- tion that Australia and New Guinea with their orig- inally poor Subantarctic flora, had drifted together from the Subantarctic regions towards the NW and so had come, in the Upper Tertiary, into contact Fig. 6. Comparison of the character of the Phi- lippine flora by contrasting A (Asiatic-centred genera of type 2) and A’ (West Malaysian-centred genera of type 3) against B (Australian and Subant- arctic genera of types 4 & 5) and B’ (East Malaysian genera of type 3). with the Malaysian tropical plant world.! The Ma- laysian vegetation, then, overwhelmed this original Subantarctic flora and its remnants could only sur- vive on the high mountains of New Guinea. This certainly contradicts the opinion of Hoo- KER, BENTHAM, and Dies who in their analyses (1) Blumea 1 (1934) 115-159; in: KLEIN, Nieuw Guinee 1 (1935) 192-198. Library oe es a = 4 = 142) (@>) vt AN © N >) York Botanica New 85 0